ISSN 0968-0454
Bulletin of
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Entomology Series
VOLUME 63 NUMBER 2 24 NOVEMBER 1994
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© The Natural History Museum, 1994
Entomology Series
ISSN 0968-0454 Vol. 63, No. 2, pp. 137-210
The Natural History Museum
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Printed in Great Britain at The Alden Press, Oxford
Bull. Br. nat. Hist. (Ent.) 63(2): 137-210
Issued 24 November 1994
A taxonomic review of the common
green lacewing genus Chrysoperla
(Neuroptera: Chrysopidae)
S.J. BROOKS
Department of Entomology, The Natural History Museum, Cromwell Road,
London SW7 5 BD
CONTENTS
Introduction 137
Historical review 138
Generic classification 1 38
Species classification 1 39
Materials and methods 140
Abbreviations 1 40
Acknowledgements 141
The systematic position of Chrysoperla 141
Species affinities within Chrysoperla 142
Chrysoperla Steinmann 143
Key to the world Chrysoperla species 144
Species descriptions 146
Synoptic checklist of the species of Chrysoperla 173
References 1 74
Index 210
Synopsis. The genus Chrysoperla Steinmann (Neuroptera: Chrysopidae), world-wide
distribution, is revised. Several species of the genus, and in particular Chrysoperla
carnea (Stephens), are cultured extensively for use in biological control programmes but
until now the systematics of the genus has remained confused. 36 species are recognized
as valid. Five new species are described, 4 species have revised status and there are 40
new specific synonyms, one new provisional synonym and four lectotype designations.
Eight species are newly excluded from Chrysoperla. A key to the species is provided.
Male and female of each species are described and diagnostic features figured.
INTRODUCTION
More effort has been spent on research into the
ecology and biology of Chrysoperla than all the
rest of the chrysopid genera put together (see
Canard et al., 1984). This is chiefly because
several species, but especially C. carnea
(Stephens), have proved easy to culture and their
larvae are fierce predators of aphids and coccids.
C. carnea and some of its congeners are now
commonly reared in biological control laborato-
revision of the
Barnard, 1990),
defined and it is
ries all over the world. One aspect that has been
largely overlooked, however, is the systematics
of the genus as a whole.
Following a genus-level
Chrysopidae (Brooks &
Chrysoperla is now clearly
possible to correctly assign species to the genus.
As currently recognized, the genus includes 36
species, distributed throughout the world but
with a preponderance of Holarctic species.
Although it is relatively easy to recognize which
species belong to the genus, species level identifi-
)The Natural History Museum. 1994
138
S.J. BROOKS
cation is less straight forward. Tjeder (1966)
provided a key to five species from southern
Africa and Holzel (1989b) gave descriptions of a
further five Afrotropical species. Three western
European species can be identified using the keys
in Aspock et al (1980). In order to identify the
remaining Old World species it is usually neces-
sary to compare the frequently inadequate origi-
nal descriptions that are scattered throughout the
literature often in old and obscure volumes. The
New World also has a rich Chrysoperla fauna but
similar identification problems will be encoun-
tered as there are no modern keys available to
identify the adults, although Tauber (1974) pro-
vided descriptions of the North American larvae.
A further complication when trying to estab-
lish the correct name for a particular Chrysoperla
specimen is that, although a few identification
keys are available for particular regions, regional
faunas have never been compared with each
other. This has undoubtedly led to synonymy and
this study shows that some species occur in more
than one zoogeographic region where they were
referred to under different names.
In recent years traditional morphological taxo-
nomic studies of Chrysoperla species, especially
those in the C. carnea species-complex, have
been supplemented by investigations employing
additional techniques. One example is the pio-
neering work begun by Henry (1979) who stud-
ied the production of substrate-borne calls by
males and females during courtship. This work
has necessitated a reappraisal of species limits
because morphologically identical populations
have been shown to be reproductively isolated by
their different courtship songs (Henry, 1983). A
useful summary of this work appears in Henry
(1992). Other new approaches include multivari-
ate analysis of morphological characters (Thierry
et al., 1992), morphometric studies of the male
genitalia (Semeria, 1992) and electrophoretic
studies (Cianchi & Bullini, 1992). These and
other studies on the Chrysoperla carnea-complex
are discussed by Thierry & Adams (1992).
These recent investigations have produced
some fascinating results and point up some of the
limitations of a traditional morphological taxo-
nomic study. However, a thorough revision of
the genus on a worldwide basis is an essential
prerequisite if we are to interpret the phyloge-
netic relationships of the species within the
genus. Such a revision will also establish the
geographic distribution of the morpho-species
and the extent of geographical variation within
taxa. The new techniques described above have
only been applied to a few populations in western
Europe and North America where a plentiful
supply of material is readily available. In order to
conduct a comprehensive revision of the world
species it is necessary to fall back on a morpho-
logical study of long-dead material stored in
museum collections.
This study is the first revision of the world
species of this economically important genus. As
a result of this work 36 species are now recog-
nized as valid and these are described and fig-
ured. Five new species are described, 4 species
have new status, 40 new specific synonyms, three
revised synonyms and one provisional synonym
are proposed. The aim of this work is to provide
an overview of the species of Chrysoperla and a
baseline for future taxonomic work, using a
wider range of techniques.
In general, I have decided to recognise as
species those taxa that have clearly defined,
unambiguous, morphological characters. Such
characters are usually derived from features of
the male genitalia but include other characters
too. Geographical variation within each species
is described. Such variation may be indicative of
sibling species that may be more clearly revealed
using some of the techniques described above. A
few species have recently been described that are
distinguishable only by the courtship songs that
they produce or by certain aspects of their ecol-
ogy. I have not synonymised these taxa since the
biological evidence is compelling and active
research is still in progress.
HISTORICAL REVIEW
Generic classification
Chrysoperla was first proposed by Steinmann
(1964) as one of four subgenera of Chrysopa
Leach. The subgenera were based on differences
in head markings. However, characters such as
these have proved to be unreliable in defining
supraspecific monophyletic groups (Brooks &
Barnard, 1990) and Steinmann's subgenera have
no validity as he defined them. The systematics
of Chrysopidae was revolutionized when Tjeder
(1966) recognized that the presence or absence of
certain components in the male genitalia united
groups of species in the otherwise homogeneous
genus Chrysopa s.l. Until then this large and
unwieldy taxon included the large majority of
chrysopid species. Tjeder considered these
groups of species as subgenera of Chrysopa. One
of the eight subgenera he recognized included
those species in which the arcuate dorsal struc-
ture in the male genitalia, termed the tignum.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
139
was present but the gonapsis, a ventral winged
structure, was absent. This species group
included C. cornea (Stephens) which Steinmann
had nominated the type-species of Chrysoperla.
Tjeder (1966) included 17 species in Chrysoperla
which were divided between two species groups.
The second of these species groups is now consid-
ered to be a distinct genus, Atlantochrysa Holzel
(Brooks & Barnard, 1990). Following Tjeder's
lead. Holzel (1970) revised the generic classifica-
tion of the Palaearctic Chrysopinae. Holzel
accorded full generic status to some of the more
distinctive taxa previously grouped together in
Chrysopa s.l. but those groups in which males
possessed a tignum and/or gonapsis, including
Chrysoperla, were included together as subgen-
era of Anisochrysa Nakahara (now Mallada
Navas).
Semeria (1977) later raised Chrysoperla to full
generic status. He showed that species of
Chrysoperla were distinct from those of Anisoch-
rysa in three principal ways. First, males of
Anisochrysa possess a gonapsis, which is absent
in Chrysoperla, secondly larvae of Anisochrysa
carry debris but are naked in Chrysoperla and
finally, the adult is the over-wintering stage in
Chrysoperla but in Anisochrysa it is the third
instar larva that over-winters.
Electrophoretic studies by Bullini et al. (1984)
supported the recognition of Chrysoperla as dis-
tinct from other chrysopid genera and moreover
demonstrated that it was more closely related to
Anisochrysa (= Mallada) than to Chrysopa
Leach.
The first genus-level revision of the world
Chrysopidae was published by Brooks & Bar-
nard (1990). In that paper, Chrysoperla was
demonstrated to be monophyletic, closely related
to but distinct from Mallada. As well as the
characters noted by previous workers as distin-
guishing Chrysoperla, additional autapomorphies
for the genus were identified.
Species classification
Although the species of Chrysoperla are superfi-
cially rather homogeneous, there is considerable
variation in the extent and colour of markings,
especially on the head, within and between taxa.
Many of the taxa are also common, abundant
and widely distributed. This combination of fac-
tors has lead to a plethora of species-group
names appearing in the literature. During this
study 112 species-group names assignable to
Chrysoperla were traced (plus numerous varietal
names) but I have treated only 36 as valid.
The wide distributional range and degree of
morphological variation between populations of
some Chrysoperla species led some authors to
doubt whether these taxa are true biological
entities. In order to tackle this problem some
workers have adopted additional methods that
supplement the traditional use of morphological
characters. These methods include the analysis of
courtship songs", multivariate analysis of mor-
phological characters, morphometric methods
and biochemical techniques.
Henry (1979) investigated courtship and mat-
ing in North American examples of C.
plorabunda and found that rhythmic tremulation
of the abdomen occurred in both sexes through-
out courtship. These non-audible vibrations arc
transmitted through the substrate and picked up
by receptors in the insects' legs. Further work
(e.g. Henry. 19X3: 1985a & b; Henry & Wells.
1990; Henry et al., 1993) has revealed the exist-
ence of song morphs' that are morphologically
indistinguishable but produce different courtship
songs. In some Nearctic localities these song
morphs occur sympatrically but will not naturally
interbreed and Henry (1993) has described these
as distinct sibling species. However, the signifi-
cance of song differences in at least some
instances remains unclear. Wells (1991) was able
to produce fertile hybrids in the laboratory
between two North American song-morphs (C.
plorabunda and ('. johnsoni) and Duelli (1987)
reported that hybridisation did occur in the labo-
ratory between two song morphs in Switzerland.
Wells (1991), using gel electrophoresis, has
found the genetic distance between three song-
morphs of C. plorabunda to be no greater than
that usually found between individuals of the
same animal species. The songs of few European
populations have been examined but Henry
(1983) has shown that individuals of C. carnea
from Switzerland produce songs that are radi-
cally different from those of otherwise indistin-
guishable nearctic populations.
Thierry el al., (1992) have adopted a different
approach and conducted a multivariate analysis
of eight morphological character states in French
populations of C. carnea. Their results indicate
that there are three distinct adult morphs that
can be associated with three larval morphs.
While refraining from naming these morphs as
distinct species, they suggest that these morphs
may represent sibling species.
Detailed morphometric studies of the male
genitalia by Semeria (1992) have suggested the
occurrence of six sibling species of Chrysoperla
carnea in France.
Electrophoretic and genetic studies of Euro-
pean populations of C. carnea have also sug-
140
S.J. BROOKS
gested the presence of sympatric sibling species
(Bullini & Cianchi, 1984; Bullini et at., 1983;
1984; Cianchi & Bullini, 1992). Results suggest
that certain central European populations are
reproductively isolated. However, the values of
genetic distance (Nei, 1972) are relatively low so
it is postulated that they have recently diverged.
The occurrence of sibling species in the C.
carnea complex has also been identified follow-
ing ecological studies. Tauber & Tauber (1977 a;
b) identified two sympatric sibling species within
Nearctic populations of the cornea-complex . One
of the sibling species, C. downesi, occurs in
conifer forests and is dark green all year round.
The second sibling species, C. plorabunda,
occurs in meadows during summer, when it is
bright green, but during the winter is found in
deciduous woodland and is brown. Morphologi-
cally these two taxa are otherwise indistinguish-
able.
The work described above presents difficulties
in interpretation especially when the genus as a
whole is being reviewed. First, only a few taxa
have been investigated in such ways and then
from only a limited geographical range. Also
there has been no integration between these
techniques. At present none of the so-called
song-morphs have been examined using multi-
variate techniques and few have been investi-
gated using electrophoresis. DNA studies of any
of the above sibling species may also produce
useful comparative results.
Until Chrysoperla populations have been
investigated more comprehensively I think that it
is unwise to introduce even more names to an
already confusing picture. In this work I have
decided to recognise species if they can be clearly
defined using classic morphological characters,
such as those that occur in the male genitalia, or
if there is overwhelming evidence from other
techniques. This should provide a sound frame-
work on which to base further studies on the
possible occurrence of sibling species.
examined and in many cases the genitalia dis-
sected. Drawings were made using a camera
lucida attachment on a stereo dissecting micro-
scope or compound microscope. The genitalia
were stored in glycerol in glass vials or perma-
nently slide mounted. The type specimens of
many species were examined but, where this was
not possible, reliably identified specimens were
examined instead.
In the synonymic lists provided with each
species description only species and subspecies
are listed. For several species large numbers of
named varieties have also been described. Most
of these have not been examined. If this had
been attempted the length of time spent on the
project would have been considerably increased
but almost certainly would have had no impact
on the validity of the species-group names used
in this study.
There are several morphological characters
which were found to be particularly useful when
distinguishing species or determining phyloge-
netic relationships. Some of these involved com-
parative measurements using a calibrated
graticule.
Length of costal setae
The length of the fringe of setae on the costal
margin of the fore wing was determined by
measuring the height of the fringe at right angles
to the costal vein at the level of the second radial
crossvein. The more erect the setae, the longer
the costal fringe will appear.
Fore wing length: breadth
This ratio is obtained by dividing the overall
length of the fore wing by the width at the wing's
widest point, measuring perpendicularly from
the costa.
Basal dilation of tarsal claw
MATERIALS AND METHODS
The relative size of the basal dilation is expressed
as a ratio of the claw hook length to the length of
the basal dilation of the hind tarsal claw.
This work has been based primarily on the exten-
sive collections of Chrysoperla specimens held at
The Natural History Museum, London, although
additional specimens have been kindly loaned by
many institutions and individuals around the
world. The types of 61 of the 117 described
species assignable to Chrysoperla have been
studied and over 2000 specimens have been
ABBREVIATIONS
BAU Beijing Agricultural University, Beijing,
China
BMNH The Natural History Museum, London, UK
BPBM Bernice P. Bishop Museum, Honolulu, USA
CAS California Academy of Sciences, San Fran-
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
141
cisco, USA
CNC Canadian National Collection, Ottawa,
Canada
HU Hokkaido University, Sapporo, Japan
IP Institut fur Pflanzenschutzforschung, Eber-
swalde, Germany
IRSM Institut de Recherche Scientifique de Mada-
gascar
IRSNB Institut Royal des Sciences Naturelles de
Belgique, Bruxelles, Belgium
ITZ Instituut voor Taxonomische Zoologie,
Amsterdam, The Netherlands
IZAS Institute of Zoology Academia Sinica,
Beijing, China
KU Kyushu University, Fukuoka. Japan
LACM Los Angeles County Museum, Los Angeles,
USA
LP La Plata Museum, Buenos Aires, Argentina
MCSN Museo Civico di Storia Naturale, Genoa,
Italy
MCZ Museum of Comparative Zoology, Cam-
bridge, USA
MNHN Museum National d'Histoire Naturelle,
Paris, France
MRAC Musee Royal de l'Afriquc Centrale, Tcr-
vuren. Belgium
NCI National Collection of Insects, Pretoria,
Republic of South Africa
NHMZ Natural History Museum of Zimbabwe.
Bulawayo, Zimbabwe
NMHU Museum fur Naturkunde der Humboldt-
Univcrsitat. Berlin, Germany
NMW Naturhistorisches Museum Wien, Vienna,
Austria
NNM Nationaal Natuurhistorisch Museum,
Leiden. The Netherlands
NR Naturhistoriska Riksmuseum. Stockholm.
Sweden
SDNHM Natural History Museum, San Diego. USA
TM Transvaal Museum, Pretoria, Republic of
South Africa
USP Museum of Zoology, University of Sao
Paulo, Brazil
ZMH Zoologisches Museum, Hamburg, Germany
ZSBS Zoologisches Sammlung des Bayerischen
Staater, Munich, Germany
ZSI Zoological Survey of India, Calcutta, India
Acknowledgements. I would like to thank the follow-
ing for the loan of specimens: Dr P. A. Adams (Fuller-
ton University, USA), Dr H.M. Andre (MRAC), Dr
R. Gaedike (IP), Dr N. Evenhuis (BPBM), Dr D.
Faulkner (SDNHM), Mrs M.J. Fitzpatrick (NHMZ),
Dr S. de Freitas (USP), Dr C.T. Graham (MCZ), Dr
P. Grootaert (IRSNB), Prof. C.S. Henry (University of
Connecticut, USA), Mr W. Hogenes (ITZ), Herr H.
Holzel (Bruckl, Austria), Dr Kuldip Rai (ZSI), M J.
Legrand (MNHN), Dr B. Gustafsson (NR), Dr M.W.
Mansell (NCI), Dr V.J. Monserrat (University of
Madrid, Spain), Prof. S. Nomura (KU), Dr N.D.
Penny (CAS). Mr C.W. Plant (Passmore Edwards
Museum. UK). Dr R. Poggi (MCSN). Mr R.R. Snel-
ling (LACM), Dr S. Takagi (HU). Prof. C.A. Taubcr
(Cornell University, USA), Mr J. van Tol (NNM), Dr
X.-k. Yang (IZAS).
I am also grateful to the following for fruitful
exchange of ideas: Dr P. A. Adams (California State
University, Fullerton, USA). Prof. C.S. Henry (Uni-
versity of Connecticutt, Storrs, USA). Prof. E.G.
MacLeod (University of Illinois, Urbana, USA), Prof.
C.A. Tauber (Cornell University. Ithaca. New York).
THE SYSTEMATIC POSITION OF
CHRYSOPERLA
Brooks & Barnard (1990) demonstrated that
Chrysoperla belonged to the Chrysopini which,
with 30 genera, is the largest tribe of Chrysopi-
dae. There are a number of adult characters that
may shed light on the phylogenetic relationships
of Chrysoperla.
In discussing the phylogenetic significance of
male genitalic components in Chrysopidae,
Brooks & Barnard (1990) concluded that posses-
sion of a tignum and gonapsis was autapomor-
phic for the tribe but plesiomorphic within the
Chrysopini. Because of the morphological uni-
formity of the tignum throughout the tribe, it is
unlikely that the tignum has evolved more than
once but may have been lost on more than one
occasion. An arcuate tignum occurs in six
chrysopid genera: Chrysopidia (subgenus Anach-
rysa) Holzel, Anomalochrysa McLachlan, Atlan-
tochrysa Holzel, Chrysoperla Steinmann,
Dichochrysa Yang & Yang and Mallada Navas.
Like Chrysoperla, Chrysopidia (Anachrysa) and
Mallada (satilota-group) do not have a gonapsis
but the gonapsis may have been lost indepen-
dently in these taxa. A gonapsis is present in the
other genera mentioned above including the
Mallada desjardinsi-group.
The yellow median thoracic band is a charac-
teristic feature of most Chrysoperla species.
Within the family this marking is restricted to the
chrysopine genera Anomalochrysa, Apertochrysa
Tjeder, Chrysoperla, Chrysopidia (subgenus
Chrysotropia) Navas, Cunctochrysa Holzel,
Dichochrysa, Mallada and Peyerimhoffina Lac-
roix.
Many species of Chrysoperla possess dorsal
striations on the arcessus. This character is prob-
ably derived in Chrysopidae since it does not
occur elsewhere in the Neuroptera. The occur-
142
S.J. BROOKS
rence of this character in other chrysopid genera
suggests that they are closely related. There are
only five other chrysopid genera, and like
Chrysoperla all are Chrysopini, that have a stri-
ated arcesuss: Apertochrysa, Atlantochrysa,
Dichochrysa, Meleoma and Suarius Navas.
Possibly the most frequently used character for
distinguishing Chrysoperla in keys is the short
intramedian cell, the apex of which does not
usually reach the first crossvein from the radial
sector. This character, although not occurring
universally in species of Chrysoperla, may be a
synapomorphy linking Chrysoperla with two
other chrysopine genera, Mallada and Peyerim-
hoffina. The character does not occur elsewhere
in the Chrysopidae.
The sister group to Chrysoperla plus the other
chrysopine genera discussed above is probably
Dichochrysa. Dichochrysa is linked to these gen-
era by several synapomorphies but also retains
some plesiomorphies, particularly in the male
genitalia, that have become modified in the other
genera. For this reason, Dichochrysa will be
considered as the outgroup when deciding on
character polarities in postulating species rela-
tionships within Chrysoperla.
SPECIES AFFINITIES WITHIN
CHRYSOPERLA
There are four distinct species groups within the
genus based on the following adult characters.
1. carnea-group. This group is the least derived
and retains many plesiomorphic chrysopine char-
acters, particularly in the male genitalia. The
male genitalia of the group lack spinellae,
gonosetae are generally few, the arcessus is dor-
sally striated, the entoprocessus are large and
crescentic, the gonarcus arch is broad and the
lateral plates are large and reniform, and the
acumen of the tignum is narrow at its apex. Most
of these characters are plesiomorphic for the
Dichochrysa-group of genera, although the large
crescentic entoprocessus are characteristic of the
cornea-species group and are probably apomor-
phic. The carnea-group can be divided into two
sub-groups.
1.1. The first sub-group includes the species
carnea (Palaearctic), furcifera, nipponensis,
oblita, sillemi (Eastern Palaeartic), iranica
(Middle East), renoni, mediterranea (Western
Palaearctic), zastrowi (Afrotropics) and
plorabunda, adamsi, johnsoni and downesi
(Nearctic). In these species, with the exception
of C. oblita, the basal dilation of the claws is
small (apomorphic for this species group).
1.2. The second sub-group comprises gallagh-
eri (Eremic) and annae (Eastern Palaearctic).
The numerous gonosetae and small, narrow
entoprocessus in the male genitalia are genus-
level apomorphies that set this species-subgroup
apart.
The following species-groups (2, 3, and 4) all
possess numerous gonosetae which is apomor-
phic within Chrysoperla. The possession of a few
gonosetae, arranged in a lateral group, is a
synapomorph that links most of the genera in the
Dichochrysa group. Species groups 2 and 3 are
linked by the synapomorphic character of broad
fore wings and long costal setae.
2. comans-group. This group comprises
comans, decaryana (both Afrotropics) and sia-
mensis (Thailand). In these species spinellae are
absent (plesiomorphic within Chrysoperla), but
the species share two characters that are apomor-
phic within the genus: the arcessus is unstriated
dorsally and the entoprocessus are very small.
However, this species-group is not well-defined
since both these apomorphies occur in other
Chrysoperla species-groups and, in addition, an
unstriated arcessus is considered to be a second-
ary character reversal. Nevertheless, the absence
of spinellae in the male genitalia of group 2
species preclude their inclusion in groups 3 and 4
since this is autapomorphic for those groups.
3. nyerina-group. Species in this group, like
group 2, also have broad fore wings and long
costal setae, which is apomorphic within
Chrysoperla. In addition, males possess spinellae
on the gonosetae. This character is synapomor-
phic for groups 3 and 4 and does not occur
elsewhere in the Chrysopidae. The nyerina-
group can be divided into two sub-groups.
3.1. The first sub-group, comprising nyerina
and plicata (both Afrotropics), is characterised
by the presence of dorsal striations on the arces-
sus (plesiomorphic in Chrysoperla) and the pos-
session of simple spinellae. The spinellae show
several character states within Chrysoperla. C.
plicata exhibits the simplest condition in which
the spinellae arise directly from the gonosaccus.
In nyerina the spinellae are modified so that each
spine is positioned on a small tubercle.
3.2. C. insulata (Reunion Island) is the only
species in this sub-group. The spinellae are more
complex than those found in sub-group 3.1,
being formed from 2-3 spines on a small
tubercle. This is the only species in the genus
which lacks a tignum in the male genitalia. This
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
143
structure has probably been secondarily lost.
4. pudiea-group. This group is characterised
by the possession of complex spinellae formed
from 5 or more spines on each tubercle. This is
the most derived condition of the spinellae within
the genus. In this group the fore wings are
narrow with short costal setae. The shape and
length of the apex of the arcessus was found to be
particularly useful in distinguishing species of this
group. There are two sub-groups.
4.1. This sub-group includes the species
pudica (Afrotropics), mutata and nigrinervis
(Palaearctic). The gonarcus retains its typical
broad arch configuration with large, reniform
lateral plates and small entoprocessus. The acu-
men is swollen apically. At present no apomor-
phic characters have been found to define this
sub-group.
4.2. In the second sub-group most species are
united by the autapomorphy of a narrow gonar-
cus arch (which does not occur elsewhere in
Chrysoperla or in related genera) with narrow
lateral plates and minute entoprocessus. How-
ever, dozieri and defreitasi are exceptional in
retaining a relatively broad gonarcus arch. Spe-
cies included are: exul (St Helena), congrua (Old
World tropics), asoralis, exotera, externa, galapa-
goensis, mexicana (all Nootropics). These species
all possess an apically swollen acumen. However,
in the species, comanche, harrisii, rufilabris (all
Nearctic), defreitsi (Brazil) and dozieri (Haiti),
the acumen is narrow apically.
Chrysoperla Steinmann
Chrysoperla Steinmann, 1964: 260 [as subgenus
of Chrysopa Leach]. Type-species: Chrysop-
erla carnea Stephens, 1836, by original desig-
nation. [As subgenus of Anisochrysa Nakahara
by Holzel, 1970: 51; raised to genus by Seme-
ria, 1977: 238.]
Adult. Fore wing 9-14 mm. Ground colour
pale green, becoming brown in over-wintering
specimens. Head sometimes unmarked but usu-
ally with red or blackish brown stripe on gena
and lateral clypeus. Maxillary palps unmarked or
with black dorsal stripe. Labrum indented anteri-
orly. Mandibles broad, asymmetrical with basal
tooth on left mandible. Head width : eye width =
1.8-2.6 : 1. Antenna usually shorter than fore
wing; flagellar segments 2-3 times as long as
broad; setae arranged in four rings. Pro-, meso-
and metanotum usually with yellow or white
longitudinal median band, sometimes marked
with lateral red spots; dorsal setae long and pale
or short and dark. Leg unmarked; claw with, or
occasionally without, basal dilation. Fore wing
unmarked; narrow or broad (length : breadth =
2.6-4.1 : 1); venation green or with gradate
crossveins black; costal setae short, inclined
towards wing apex or sometimes long, erect;
basal costal crossveins sinuous; stigma
unmarked; cell im narrow, ovate; 1st Rs cross-
vein usually meeting Psm distad of apex of im,
but sometimes meets im at apex or subapically;
gradates in two parallel series, basal inner gra-
date meets Psm. Abdomen usually with longitu-
dinal median yellow or white stripe.
GenitaliacT. Apex of sternite 8+9 with distinct
lip beset with short setae. Ectoproct with invagi-
nation ventrad of callus cercus. Gonosaccus with
gonosetac. with or without spinellae. Arcessus
narrow with or without dorsal striations. Ento-
processus crescentic, large or minute. Gonarcus
broadly arcuate with reniform lateral plates or
arch narrow with narrow lateral plates. Tignum
arcuate; acumen long and narrow or short, swol-
len. Gonapsis absent.
GENITALIAS'. Praegenitale absent. Subgenitale
bilobed apically. tapering and extended basally.
Spermatheca narrow; ventral impression slight:
vela short; duct short.
Larva. Abdomen narrow. fusiform. not
humped. Thoracic and abdominal tubercles
small, spherical; setae short, smooth, not hooked
apically: transverse row of metanotal setae
absent; latero-dorsal chalazae present but indis-
tinct, bearing single seta. Debris not carried.
Rt marks Adult males of Chrysoperla are char-
acterised by the autapomorphic possession of a
lip at the apex of sternite 8+9. The fusiform
larvae with small tubercles and non-debris carry-
ing habit are also distinctive. These characters
are apomorphic at the genus-level. Other auta-
pomorphies, but which are not shared by all
species in the genus, include: the presence of
spinellae and an over-wintering diapause occur-
ring in the adult stage.
As a result of this study it is now apparent that,
in order to maintain Chrysoperla as a monophyl-
etic group, a few species listed under Chrysoperla
by Brooks & Barnard (1990) should be moved to
different genera. The confusion has arisen
because the possession of a tignum but lack of a
gonapsis was previously considered apomorphic
for Chrysoperla. However, it is now clear that
the gonapsis has been lost independently at least
twice in the Chrysopidae. Consequently, the
species anpingensis Esben-Petersen, australis
New, krakatauensis Tsukaguchi, maquilingi
144
S.J. BROOKS
Banks, meloui Navas, satilota Banks, suzukii
Okamoto and triactinata New, in which sternite
8+9 is not lipped in males, must be reassigned
(summarised in checklist, p. 173).
Yang & Yang (1990b) proposed that the Mal-
lada boninensis (Okamoto) (now a synonym of
M. desjardinsi Navas (Holzel & Ohm, 1992))
-group of species (Brooks & Barnard, 1990) be
considered as a distinct genus and that the
remaining Mallada species be included in a new
genus, Dichochrysa Yang & Yang. This proposal
has merit since there are several autapomorphies
that define Mallada s. str., and Mallada s.l.
seems to be paraphyletic.
The males of Mallada s. str. are characterised
by narrow ectoprocts, subapical lobe on sternite
8+9, two pairs of gonosetae in the genitalia and
in the fore wing by the basal inner gradate
meeting Psm. Species in the desjardinsi-group
possess a tignum with a minute acumen, a broad
arcessus and a cruciform, basally expanded
gonapsis. The following species are transferred
to Mallada and are members of the desjardinsi-
group: anpingensis Esben-Petersen comb, n.,
meloui Navas comb. n. and maquilingi Banks ( =
inconspicua Navas) comb. n. In a second species
group within Mallada s. str. the gonapsis is
absent (instead there is a large V-shaped apo-
deme at the apex of sternite 8+9), the gonosetae
are simple (in the desjardinsi-group the setal
bases are thickened so that they resemble
mounted needles), the arcessus is narrow and
elongate and the acumen is short and ovate. It is
now clear that some species formerly included in
Chrysoperla belong to this second species-group
and so are reassigned to Mallada: krakatauensis
Tsukaguchi comb, n., satilota Banks comb. n.
and suzukii Okamoto comb. n.
The presence of gonocristae in australis and
triactinata suggest a relationship with Aperto-
chrysa, although species of this genus lack a
tignum and possess a gonapsis. It may be neces-
sary to erect a new genus for these two Austra-
lian species but for the present they are
provisionally assigned to Apertochrysa: Aperto-
chrysa australis comb, n., Apertochrysa triacti-
nata comb. n.
I have been unable to examine the types of the
following species that are deposited in BAU or
IZAS, China: Chrysoperla quinlingensis Yang &
Yang, C. thelephora Yang & Yang, C. yulinica
Yang & Yang (Yang & Yang 1989), C. bellatula
Yang & Yang, C. hainanica Yang & Yang, C.
longicaudata Yang & Yang, C. sola Yang &
Yang (Yang & Yang, 1992) or Chrysopa xizan-
gana Yang (Yang, 1988). Unfortunately, it is not
possible to decide the affinitites of these species
from the original descriptions so at present they
must remain incertae sedis.
Distribution. Eight species are known from
the Afrotropics, six from the Western Palaearctic
region, three from the Eastern Palaearctic, there
is one Palaearctic species, three Oriental species,
one species occurs throughout the Old World
tropics, there are ten Nearctic species and four
Neotropical species.
Key to the world Chrysoperla species
This key does not include the following Chinese
species which were unavailable for examination:
Chrysoperla quinlingensis Yang & Yang, C.
thelephora Yang & Yang, C. yulinica Yang &
Yang, C. bellatula Yang & Yang, C. hainanica
Yang & Yang, C. longicaudata Yang & Yang, C.
sola Yang & Yang or Chrysopa xizangana Yang.
1 Gradates green 2
— Gradates black or greyish 17
2 Base of claw undilated or only slightly swollen (Figs
33.51) 3
— Base of claw with distinct step-like dilation, at least
one-quarter length of claw hook (Fig. 5) 8
3 Head with extensive black/brown markings, with
X-mark between antennae furcifera Okamoto
— Markings on head restricted to red or brown stripe
on gena. frons or clypeus 4
4 Antennae black/brown; veins 1A and 2 A of fore
wing black/brown mexicana sp. n.
— Antennae green; veins 1A and 2A of fore wing
green 5
5 Intramedian cell (im) long, apex of im meets first Rs
crossvein (Fig. 32) 6
— Intramedian cell short, apex of im not reaching first
Rs crossvein (Fig. 1) 7
6 Post-ocular region marked red; gena marked with
red spot; apex of fore wing pointed; cf genitalia
with spinellae harrisii Fitch
— Post-ocular region unmarked; gena marked with
brown spot; apex of fore wing rounded; spinellae
absent in c$ genitalia annae sp. n.
7 Costal setae long, 1.5 mm; wing apex rounded (Fig.
68); claw hook very long (Fig. 69) . renoni Lacroix
— Costal setae short mediterranea Holzel
8 Basal dilation of claw large, at least half length of
claw hook
— Basal dilation of claw small, distinctly less than half
length of claw hook 10
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
145
9 Pronotum marked with brown spot in each corner
in addition to yellow median band; vertex marked
with red lateral spot exul McLachlan
— Pronotal markings restricted to yellow median
band; vertex unmarked comanche Banks
10 Gena usually marked with red spot; red post-ocular
spot usually present; cf: spinellae present 11
— Gena usually marked with brownish/black spot,
which may be overlaid with red; post-ocular region
usually unmarked; cf: spinellae absent 12
1 1 Basal dilation of claw less than third length of claw
hook (Fig. 189); Old World tropics cf: neck of
acumen of tignum short, swollen apically
congrua Walker
— Basal dilation of claw about third length of claw
hook (Fig. 228); Neotropics; cf: neck of acumen
long, swollen apically externa Hagcn
12 Fore wing long, more than 13 mm; cf: 10-20
gonosetae lateral group annae sp. n.
— Fore wing shorter, usually less than 13 mm; cf: few
gonosetae (less than 10 in lateral group) 13
13 Fore wing rounded, symmetrical apically (Fig. 40);
Cf: membranous lip of sternite 8 + 9 narrow, elon-
gate (Fig. 42) iranica Holzel
— Fore wing tapered, asymmetrical apically (Fig. 3)
(in some northern European specimens fore wing
may be rounded with long setae Fig. 2); cf: mem-
branous lip of sternite 8+9 short, broad, rounded
(Fig. 8) 14
14 Basal dilation of claw one-quarter or less length of
claw hook (Fig. 25) 15
— Basal dilation of claw more than third length of claw
hook 16
15 First Rs crossvein at oblique angle to Rs; Afrotro-
pics zastrowi Esben-Petersen
— First Rs crossvein usually at right angles to Rs;
India sillemi Esben-Petersen
— First Rs crossvein variable; Nearctic
19 Cf: spinella present
20
downesi Smith
16 Palaearctic carnea Stephens
— Nearctic plorabunda Fitch species group
17 Gena marked with red spot or stripe; if unmarked,
clypeus with black spot [some specimens of the
plorabunda species-group complex may key out at
this point] 18
— Gena marked brown/black, often with red overlay;
if unmarked, clypeus with red spot 30
18 Costal setae of fore wing long, at least 1 mm, often
erect 19
— Cf: spinella absent 21
20 Gena marked with red spot, clypeus unmarked:
pronotum marked with red stripe in each corner
and red medio-lateral spot; claw rounded posteri-
orly, with very large basal dilation (Fig. 134); fore
wing less than 13 mm; cf: apex of sternite 8+9
narrow, prominent (Fig. 135); acumen of tignum
short, swollen apically (Fig. 136) ... plicata Tjeder
— Gena and clypeus marked with red stripe; any red
spots on pronotum restricted to anterior corner;
claw straight posteriorly with smaller basal dilation
(Fig. 125); fore wing more than 13 mm; cf: apex of
sternite 8 + 9 short and broad (Fig. 126); acumen
long, narrow apically (Fig. 127) .... nyerina Navas
21 cf: arcessus striated dorsally (Fig. 79)
oblita Holzel
— O": arcessus without dorsal striations 22
22 Fore wing less than 12 mm; cf: arcessus narrow
(Fig. 104); acumen of tignum short, swollen (Fig.
105); apex of sternite 8 + 9 prominent, longer than
membranous lip (Fig. 101) comans Tjeder
— Fore wing more than 12 mm; cf: arcessus broad
(Fig. 113); acumen long, parallel-sided (Fig. 112);
sternite 8 + 9 not elongated apically (Fig.
Ill) decaryana Navas
23 Maxillary palps with black dorsal stripe 24
— Maxillary palps unmarked 27
24 Red post-ocular spot present 26
— Post-ocular region unmarked 25
25 cf: apical membrane of arcessus narrow, elongate
(Fig. 281); acumen of tignum short, broad, swollen
apically (Fig. 280) rufdabris Burmeister
— Cf: apical membrane of arcessus short, broad,
tapered at apex (Fig. 216); acumen long, swollen
apically (Fig. 215) asoralis Banks
26 Front of vertex marked with two narrow red spots;
Cf: arcessus abruptly widened at apex; gonarcus
arch broad (Fig. 202) dozieri Smith
— Front of vertex unmarked, but Cuban specimens
may have a pair of broad red spots; cf: arcessus
narrow apically: gonarcus arch narrow (Fig.
22 1 ) exotera Navas
— Front of vertex unmarked; cf: arcessus broad,
narrowed subapically, spatulate at apex (Fig.
206) defreitasi sp. n.
27 Basal dilation of claw less than half length of claw
hook 28
— Basal dilation of claw more than half length of claw
hook 29
Costal setae short, inclined towards wing apex . 23 28 Basal dilation of claw about one-quarter length of
146
claw hook; scape with red stripe cf: spinellae
absent gallagheri Holzel
— Basal dilation of claw distinctly less than one-
quarter length of claw hook; scape unmarked; cf :
spinellae present nigrinervis sp. n.
29 Basal dilation of claw less than two-thirds length of
claw hook mutata McLachlan
— Basal dilation of claw hook about two-thirds length
of claw hook pudica Navas
30 Gena unmarked; intramedian cell (im) long,
extends beyond first Rs crossvein (Fig. 115) .... 31
— Gena marked with brown spot; im short, at most Rs
crossvein meets apex of im (Fig. 90) 32
31 Palps marked with black dorsal stripe; black lateral
stripe on clypeus; post-ocular region unmarked; cf :
tignum absent; spinellae present ... insulata Fraser
— Palps unmarked; red spot on clypeus; red post-
ocular spot present; cf: tignum present; spinellae
absent siamensis sp. n.
32 Basal dilation of claw less than half length of claw
hook (Fig. 238) ; crossveins in basal half of fore wing
of fore wing entirely black/brown; pronotum with
narrow black median stripe; cf: spinellae present-
galapagoensis Banks
— Basal dilation about half as long as claw hook;
crossveins in basal half of fore wing black/brown at
each end, middle section green; pronotum with
yellow median band; cf: spinellae absent 33
33 USA johnsoni Henry, Wells & Pupedis
— Eastern Palaearctic nipponensis Okamoto
SPECIES DESCRIPTIONS
Chrysoperla carnea (Stephens)
(Figs 1-4, 6-13, 17, 19-20)
Chrysopa carnea Stephens, 1836: 103. Lecto-
type?; ENGLAND: near London (BMNH)
[examined]. [Lectotype designated by Leraut,
1991: 78.]
Chrysopa affinis Stephens, 1836: 104. Lecto-
type?, ENGLAND: near London (BMNH)
[Lectotype designation by Leraut, 1991: 78.]
[examined]. [Synonymized by Schneider, 1851:
68.]
Chrysopa microcephala Brauer, 1850: 6. Holo-
type, AUSTRIA: Vienna (NMW) [not exam-
ined]. [Synonymized by Navas, 1908: 13.]
Chrysopa vulgaris Schneider, 1851: 68. Syntypes,
Europe [not traced]. [Synonymized by Killing-
ton, 1931: 112.]
S.J. BROOKS
Chrysopa lamproptera Stein, 1863: 419. Syn-
types, YUGOSLAVIA: Dalmatia, Casteln-
uovo (Kruper) (NMHU) [not examined].
[Synonymized by Aspock et al., 1980: 268.]
Chrysopa lucasina Lacroix, 1912: 203.
Lectotypecf, ALGERIA: Frenda, 1911
(Lucas) (MNHN) [Lectotype designated by
Leraut, 1991: 78.] [examined]. [Synonymized
with carnea Stephens by Aspock et al., 1980:
268; reinstated as valid species by Leraut,
1991: 76.] Syn. n.
Cintameva angelnina Navas, 1931a: 84. Holo-
type, GERMANY: Satrup (Angeln), Anfang,
ix.1930 (Lichtwerk) (ZMH) [type destroyed].
[Nomen dubium; synonymised by Aspock et
al., 1980:410.]
Chrysopa ferganica Navas, 1933b: 107. Syntypes,
USSR: Turkestan, Fergana, Sphara,
25.vii.1928 (NMHU) [not examined]. [Synony-
mized by Aspock etal., 1980: 268.]
Chrysopa pictavica Lacroix, 1933: 147. Lecto-
typecf; FRANCE: Deux-Sevres, marais
d'Amure, 27. vi. 1921 {Lacroix) (MNHN) [Lec-
totype designated by Leraut, 1982: 48.] [exam-
ined]. [Synonymized by Aspock et al., 1980:
268.]
Chrysopa carnea var. adaptata Navas, 1934a: 1.
Holotype, CHINA: Jiangsu Prov., Hufu
(Loubou), 3.viii. 1933 (IZAS) [not examined].
[As subspecies of nipponensis (Okamoto)
Yang & Yang, 1990a: 79.] Syn. n.
Chrysopa lundbladiTjeder, 1939: 8. Holotypecf ,
MADEIRA: Caramujo, 1250m (Lundblad)
(NRS) [not examined]. [Synonymized by
Aspock etal., 1980:268.]
Chrysopa maderensis Tjeder, 1939: 9. Holo-
typecf, MADEIRA: Rabacal, 1080m,
17.viM.viii.1935 {Lundblad) (NR) [not exam-
ined]. [Synonymized with carnea Stephens by
Aspock et al., 1980: 268; synonymized with
lucasina Lacroix by Leraut, 1991: 78.] Syn. n.
Chrysopa canariensis Tjeder, 1939: 31. Holotype
Cf , CANARY IS: Gran Canary, Tafira {Frey)
(NRS) [not examined]. [Synonymized by
Aspock et al., 1980:268.]
Chrysopa shansiensis Kuwayama, 1962: 9. Holo-
type $, CHINA: Shansi, Wangmaochen,
14. v. 1942 {Yasumatsu) (KU) [examined]. Syn.
n.
Chrysopa (Chrysoperla) carnea Stephens; Stein-
mann, 1964: 260.
Chrysopa (Chrysoperla) canariensis Tjeder;
Tjeder, 1966: 391.
Chrysopa (Chrysoperla) lundbladi Tjeder;
Tjeder, 1966: 391.
Anisochrysa (Chrysoperla) carnea (Stephens);
Holzel, 1970: 51.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
147
Chrysoperla carnea (Stephens); Canard & Lau-
deho, 1978:71.
Chrysopa carnea nanceiensis Semeria, 1980: 29.
Syntypes, FRANCE: Nancy (Poivre)
(NANCY UNIV.) [not examined]. [Synony-
mized by Leraut, 1991: 80; reinstated as valid
species by Semeria, 1992: 337.] Syn. n.
Chrysoperla shansiensis (Kuwayama); Brooks &
Barnard, 1990: 271.
Fore wing (Fig. 1) d\ 9-13 mm; 9- 11-14 mm.
Head marked with brown/black stripe on gena
and lateral clypeus, often overlaid with red
stripe; post-ocular spot absent. Maxillary palps
unmarked or with black stripe dorsally. Anten-
nae shorter than fore wing. Pronotum marked
with yellow/white longitudinal median stripe,
greyish spot sometimes present in anterior cor-
ner; prothoracic setae short, dark or sometimes
pale. Claw (Fig. 4) with small basal dilation,
about one-third length of claw hook (2.86-3.0: 1 ;
mean 2.70). Fore wing variable in width (length:
breadth = 2.7-3.1: 1); pointed or rounded api-
cally; venation green but costal and Rs crossveins
often with black spot at Sc and R respectively
and crossveins im-Psm, cl-c2, Rs-im sometimes
entirely black; costal setae short or relatively
long (0.4-1.6 mm), inclined towards wing apex;
cell im short, not reaching 1st Rs crossvein or
sometimes meeting at apex; 1st Rs crossvein
meets Rs obliquely or at 90°; 4-7 inner gradates,
5-8 outer gradates.
Genitaliao" (Figs 8-10). Lip of sternite 8 + 9
broad, rounded. Apodeme of tergite 9 long,
extending well-beyond callus cercus; callus cer-
cus broad, ovate; 23-33 trichobothria. Spinellae
absent. Gonosetae few, 4-6 in lateral clump,
often asymmetrical in number. Arcessus striated
dorsally; tip blunt; apex straight or curved ven-
trally at 45-90°. Entoprocessus large, crescentic.
Gonarcus with large, reniform lateral plates;
arch broad. Acumen of tignum long, rounded,
not swollen, apically.
GENITALIA9 (Figs 11-13). Callus cercus ovate;
37 trichobothria. Subgenitale with short basal
extension. Spermatheca with deep ventral
impression; vela long; duct long.
Remarks. The male genitalia of carnea are
practically indistinguishable from zastrowi, medi-
terranean furcifera, oblita, nipponensis and
plorabunda and these species are obviously
closely related to each other. However, carnea
can usually be differentiated from these species
by the combination of green gradate crossveins
and relatively broad wings, small basal dilation of
the claws and short, black prothoracic setae.
Chrysopa shansiensis Kuwayama is treated as a
synonym of carnea for the first time in this work.
Unfortunately, all that remains of the holotype
of shansiensis is the extreme apex of the abdo-
men (sternite 8, the ectoprocts and spermatheca)
and from this it is impossible to be certain of the
true identity of the species. In his description of
shansiensis, Kuwayama (1962) compares the spe-
cies with Nineta flava (Scopoli) and N. vittata
(Wesmael). However, the wing venation shown
in the photograph of the complete holotype of
shansiensis (Kuwayama, 1962) does not resemble
species of Nineta. Rather, the specimen appears
to be a Chrysoperla species and the description
suggests that it is a weakly marked example of
carnea.
Yang & Yang (1990a) considered C. carnea
var. adaptata Navas to be a subspecies of nippon-
ensis. They based their decision on similarities in
the female genitalia between nipponensis and
adaptata without indicating what those similari-
ties were. I have found the female genitalia of
nipponensis and carnea to be indistinguishable.
One character that readily serves to distinguish
carnea and nipponensis. however, is that the
gradate crossveins are green in carnea but black
in nipponensis. The gradates of adaptata are
green so it seems logical to treat this taxon as a
synonym of carnea.
Geographical variation. Mediterranean. A
series of specimens examined from Crete, Sar-
dinia, central Greece (Amfissa. Fokidhos). the
Peloponnese and Spain have distinctive mark-
ings. The most striking features, which distin-
guish this population from typical carnea, are the
very pale green overall ground colour of the body
and a pair of conspicuous black dorso-lateral
spots on the basal segments of the abdomen. In
addition, the head lacks any red markings and
the longitudinal median stripe on the pronotum
and abdomen is white. All crossveins are black at
each end, rather than being entirely green, the
wings may be pointed or rounded apically with
short or long costal setae respectively. The basal
dilation of the claw is larger than typical for
carnea (Fig. 6), being only slightly under half the
length of the claw hook (2.06-2.64: 1: mean
2.42).
Israel. A single male specimen from Lake
Huleh on the River Jordan was examined that
differed in several respects from typical carnea,
although the genitalia were indistinguishable
from typical carnea. Head suffused entirely red;
gena marked with a faint brown stripe; claw with
large basal dilation (Fig. 7) (2.41: 1); costal setae
148
S.J. BROOKS
long, semi-erect; fore wing apex rounded. How-
ever, the lack of further material precludes the
description of this specimen as a distinct taxon.
Europe. Leraut (1991; 1992) drew attention to
the co-occurrence of three forms of carnea in
Europe. The first, which Leraut established as
carnea s.str., has a dark brown to black stripe on
the gena, rounded apex to the fore wing (Fig. 2)
and relatively long wing setae. This form is
dominant in northern European populations but
occurs throughout the distributional range. A
second form was referred to as carnea B (Leraut,
1991) but was later elevated to species, and
identified as C. kolthoffi (Navas) (Leraut, 1992).
This taxon was said to differ from carnea s. str. in
having short, dark setae on the wing veins and
narrow arms in the tignum of the male genitalia.
However, I was unable to detect any difference
between specimens identified by Leraut as
kolthoffi and those identified as carnea. The
narrow arms of the tignum shown by Leraut
(1991) could be due to individual variation or
distortion, as is often the case in teneral or
immature specimens. In any case, C. kolthoffi is
an incorrect name for this taxon since kolthoffi is
a synonym of the Eastern Palaearctic species,
Chrysoperla nipponensis (Okamoto). The third
form, which Leraut (1991) referred to as C.
lucasina (Lacroix), differs from C. carnea in
having a paler stripe on the gena, short costal
setae and a straighter apico-posterior margin in
the fore wing which makes the wing appear more
pointed (Fig. 3). This form is dominant in south-
ern Europe, north Africa and the Middle East.
However, the morphological differences indi-
cated by Leraut are not as clear-cut as he sug-
gests. First, I have found every stage of genal
marking from red to dark brown on specimens
that would otherwise be referrable to lucasina
and, conversely, carnea-type specimens with a
pale brown stripe on the gena. Secondly, I have
examined several different contemporaneous
and sympatric series from various European
localities that show a gradation from the pointed
wing form with short setae to the rounded wing
form with long setae. Other specimens were
examined in which the fore wing was rounded
but costal setae short and, conversely, specimens
in which the fore wing was pointed apically and
the costal setae long. In addition, I examined
several specimens that exhibited an intermediate
condition and it was not possible to decide
whether the wings were pointed or rounded
apically.
In North America there is similar variation in
specimens of C. plorabunda with some speci-
mens having rounded wing apices with long cos-
tal setae but the majority having more pointed
wings and short costal setae. No distinction has
yet been found in the courtship songs of round
wing and pointed wing forms of C. plorabunda.
No significant difference was found in the size
of the basal dilation of the claw in specimens
referrable to f. typica or f. lucasina.
At present, I do not consider the morphologi-
cal evidence conclusive enough to treat C. luca-
sina and C. carnea as distinct species. However,
further taxonomic and biological investigations
will help to clarify this problem.
As an interesting addition to these consider-
ations, Barnard et al. (1986) reported the sympa-
tric occurrence of two forms of carnea larvae,
which had different head markings, on oak trees
in southeast England. However, although a long
series of adults was also collected throughout the
same year they were all f. typica with rounded
wing apices and long wing setae.
Using a multivariate statistical method Thierry
et al. (1992) discriminated three morphs of car-
nea in populations in central France which they
referred to as carnea, lucasina and kolthoffi. The
three main variables used to produce their results
were the colour and length of the costal setae on
the fore wing, the colour and distribution of
prothoracic setae, and the shape of the spot on
the stipe. These interesting results point the way
to further research that may establish the pres-
ence of sibling species among populations of
carnea but at present it is premature to recognise
these taxa as valid species. It is beyond the scope
of this study to conduct extensive multivariate
analysis of Chrysoperla species.
Semeria (1992) carried out a morphometric
analysis of the male genitalia of some French
specimens in the carnea species group. Essen-
tially he examined the size of the lateral plates of
the gonarcus. This was done by measuring the
angles between lines drawn from the apex of the
arcessus to the apex of the lateral plates (BAC)
and the angle between lines from the apex of the
lateral plates to the mid-point of line drawn along
the base of the lateral plates (BDC). Semeria
considered that the results indicated the presence
of six distinct species (C. renoni, lucasina, medi-
terranea, carnea, nanciensis and carnea sp. B) in
France. However, I carried out a similar analysis
of carnea group specimens and produced results
that differed widely from those of Semeria
(1992).
Because Semeria does not record any variation
in the angles he measured I assume that the
results he obtained for each taxon were derived
from only one specimen. My results show consid-
erable variation between specimens and overlap
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
149
the range measurements given by Semeria for all
the taxa he investigated. I consider it highly
unlikely that each one of the specimens I exam-
ined is a distinct species in its own right and is
also different to the taxa investigated by Seme-
ria. While the technique advocated by Semeria is
worthy of further investigation it is desirable that
a large number of specimens are examined
before any taxonomic decisions can be based on
the results. Consideration should also be given to
factors that my affect the results such as the
effects of compressing the genitalia on a micro-
scope slide. The maturity of the specimen may
also effect the results. Adams & Penny (1987)
have suggested that the genitalia may become
more heavily sclerotised as the insect matures.
This implies that the lateral plates of the gonar-
cus may actually increase in size as an individual
ages.
The morphology of the tignum shows consider-
able variation throughout the range of C. carnea
(Figs 17-20) both in length of the arms and
length of the acumen. Although I could not
detect any obvious pattern in this variation this
may merit further investigation.
Henry (1985a) reports the occurrence of two
distinct song morphs of carnea in Europe. One,
'Basel carnea\ from lowland meadows and oak
scrub in France, Switzerland and south to
Morocco. A second morph, "Ticino carnea',
occurs in the southern Swiss Alps. These taxa
cannot be distinguished from one another mor-
phologically.
Distribution. Palaearctic.
Material examined (429d\ 6959) Europe:
Belgium, Corsica, Crete, Cyprus, Czechoslova-
kia, Denmark, France, Germany, Italy, Norway,
Switzerland, Portugal, Sardinia, Spain, UK,
Yugoslavia; North Africa: Algeria, Canary Is,
Egypt, Libya, Madeira, Tunisia; Middle East:
Iran. Iraq, Jordan, Saudi Arabia, Turkestan,
Turkey, Yemen; India (Kashmir); China.
Ohm & Holzel (1992) record C. carnea from
Sudan and Senegal but all specimens examined in
this study from the Sahel region of Africa were
referrable to C. zastrowi.
Chrysoperla sillemi (Esben-Petersen) stat.
rev.
(Figs 5, 8)
Chrysopa sillemi Esben-Petersen, 1935: 234.
Holotype °- INDIA: Kashmir, Karakorum,
Maralbashi, 110m, 20.iii.1930 (Sillem) (ITZ)
[examined]. Stat. rev.
Chrysopa (Chrysoperla) gujaratensis Ghosh,
1976a: 74. Holotype o\ INDIA: Gujarat,
Dwarka, E. of Rly Stn, 2.xi.l973 (Gupta)
(ZSI) [not examined]. Syn. n.
Chrysopa (Chrysoperla) punensis Ghosh, 1976b:
189. Holotype d\ INDIA: Maharashtra,
Khed, R. Bhama, Pune, 3.vii.l976 (Ghosh)
(ZSI) [not examined, paratype examined].
Syn. n.
Chrysopa (Chrysoperla) sanandensis Ghosh,
1977: 103. Holotype d". INDIA: Gujarat,
Dadagram, Sanand, 8.xii. 1973 (Cherian) (ZSI)
[not examined, paratype examined]. Syn. n.
Fore wing 0", 10-13 mm; 9- U-14 mm. Head
occasionally unmarked or with pale brown stripe
on gena and clypeus, overlaid with red stripe.
Maxillary palps brown dorsally. Antennae
shorter than fore wing. Pronotum with yellow or
white median stripe; setae short (0.6 mm), dark
or pale. Claw (Fig. 5) with small basal dilation
3.60-5.13: 1 (mean 4.17). Fore wing length;
breadth = 2.7-3.3: 1; venation green; apex
pointed; costal setae short, inclined towards wing
apex.
GENITALIA d* and 9 as Ior C. carnea except in
male genitalia, acumen of tignum short (Fig. 8).
REMARKS. C. sillemi can be reliably distin-
guished from C. carnea only by differences in the
size of the basal dilation of the claw. In C. sillemi
the basal dilation is about one quarter the length
of the claw tooth but in C. carnea the basal
dilation is at least one third the length of the claw
tooth.
Because the ZSI has a policy of not lending
holotypes I was unable to borrow the types of the
Chrysoperla species described by Ghosh. How-
ever, paratypes were available for C. punensis
and sanandensis. These specimens were typical
examples of C. sillemi. Unfortunately, no
paratypes were available for C. gujaratensis.
Nevertheless, from the good description and
figures provided for this species (Ghosh, 1976a)
there can be little doubt that it is synonymous
with C. sillemi. The setae shown on the tignum
probably originate from the subanale.
Distribution. India, Pakistan. Specimens of
both C. sillemi and C. carnea were examined
from Kashmir.
Material examined (23d", 309) India, Paki-
stan.
150
S.J. BROOKS
Chrysoperla plorabunda (Fitch)
(Figs 15, 21-22)
Chrysopa plorabunda Fitch, 1855: 792. Syntypes,
USA: New York State & Illinois (MCZ) [not
examined]. [Synonymized with carnea
Stephens by Banks, 1903: 155; reinstated as
valid species by Henry, 1983: 299.]
Chrysopa robertsonii Fitch, 1855: 792. Syntypes,
USA:Arkansas, Tullehassie (MCZ) [not
examined]. [Synonymized by Banks, 1903:
162.]
Chrysopa pseudographa Fitch, 1855: 793. Syn-
types, USA: Illinois (MCZ) [not examined].
[Synonymized by Banks, 1903: 162.]
Chrysopa dlinoiensis Shimer, 1865: 208. USA:
Illinois[not examined]. [Synonymized by
Banks, 1903: 155.]
Chrysopa californica Coquillet, 1890: 288. Holo-
type, USA [holotype not traced]. [Smith, 1932:
594 as synonym of plorabunda; Adams, 1956:
45 as subspecies of plorabunda.] Syn. n.
Remarks. Morphologically Chrysoperla
plorabunda is indistinguishable from C. carnea.
However, Henry (1983; 1985a; 1985b and in litt.)
has provided evidence that these two taxa should
be regarded as distinct species. The substrate-
borne songs of European populations of carnea
are completely different from those of eastern
and western New World populations of
plorabunda. Egg production resulting from
crosses of the Old and New World populations of
these two taxa was very low. Although Henry's
data is derived from relatively few populations it
is persuasive and I am reluctant to synonymise
the species solely on morphological grounds. The
accrual of more song data from European and
American populations will clarify the issue.
Like carnea, specimens of plorabunda varied
in the length of the arms and acumen of the
tignum. In addition, the number of gonosetae
varied from four to eight in each lateral group.
This is a greater range of variation than that
noted in carnea. The basal dilation of the claw of
C. plorabunda is similar in size to that of C.
carnea (2.4-3.09: 1; mean 2.83)
Distribution. Nearctic.
Material examined (22cf, 38$) Canada,
Mexico, USA (Arizona, California, N. & S.
Carolina, Colorado, Florida, Montana,
Nebraska, Oregon, Texas, Utah, Vermont,
Washington, Wisconsin).
Chrysoperla johnsoni Henry, Wells &
Pupedis
(Fig 23)
Chrysoperla johnsoni Henry et al., 1993:10.
Holotype d\ USA: Oregon, Philomath
(Mary's Peak), 28.ix.1989 (Henry & Wells)
[not examined].
Fore wing d\ 10-11.5 mm; $, 11-13.5 mm.
Head marked with red stripe on gena, sometimes
with additional brown stripe; red lateral stripe on
frons and clypeus; red post-ocular spot present or
absent. Maxillary palps marked brown dorsally.
Antenna shorter than fore wing. Pronotum with
yellow median longitudinal band and greyish
lateral stripe, sometimes with reddish brown
mid-lateral spot; prothoracic setae short, pale or
dark. Claw (Fig. 15) with large basal dilation,
about half length of claw hook (2.10-2.82: 1:
mean 2.34). Fore wing narrow or broad (length :
breadth = 2.9-3.4 : 1), rounded apically; costal
setae short (0.6-0.8 mm), inclined towards wing
apex; 1st Rs crossvein meets Rs obliquely; cell im
short, not meeting 1st Rs crossvein; 4—6 inner
gradates, 5-7 outer gradates; gradates green or
black; crossveins green or black at each end
(including at wing margins); R marked with short
black stripe basal of tympanal organ. Abdomen
unmarked or marked with brown dorso-lateral
spot on segment two.
Genitalia d" and 9 indistinguishable from
Chrysoperla carnea.
REMARKS Chrysoperla johnsoni was distin-
guished from C. adamsi and C. plorabunda on
the basis of differences in their courtship songs
(Henry et al., 1993; Henry, 1993). There are two
morphs of C. johnsoni. The typical form, which
is indistinguishable morphologically from C.
plorabunda, has green gradates and the basal
dilation of the claw is about 2.8:1. The second
form, at present known only from the San Fran-
cisco Bay area, has black gradate crossveins, the
basal dilation of the claw is usually broad (about
2.2:1), the head has extensive red markings and
the gena often lacks a brown stripe. This form is
morphologically indistinguishable from the
"mohave' morph of C. downesi.
Distribution. Western USA
Material examined (7o\ 15$) USA: Ari-
zona, California.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
151
Chrysoperla adamsi Henry, Wells &
Pupedis
Chrysoperla adamsi Henry et al., 1993: 9. Holo-
type, 0\ USA: Oregon, Philomath (Mary's
Peak), 28. ix. 1989 (Henry & Wells) [not exam-
ined].
Fore wing 12-13 mm. Head marked with reddish
brown spot on gena; red stripe on gena, lateral
frons, lateral clypeus. Maxillary palps marked
black dorsally. Pronotum and vertex of head with
yellowish white median stripe; setae short, black.
Basal dilation of claws less than half length of
claw hook (2.54: 1). Wing venation green; fore
wing relatively narrow (length: breadth =
2.9-3.2: 1); apex pointed; costal setae short
(0.4-0.6 mm), inclined towards wing apex; im
short, not reaching 1st Rs crossvein; 6 inner
gradates, 6-7 outer gradates.
Genitalia o\ 9 as described for C. carnea.
Remarks. C. adamsi cannot be separated mor-
phologically from C. plorabunda, C. johnsoni or
C. downesi. However, Henry et al. (1993) has
distinguished this taxon on the basis of its court-
ship song. Unlike C. johnsoni, which may have
black gradate crossveins, specimens of C. adamsi
have green wing venation.
Distribution. Western USA
Material examined (lo\ 1$) USA: Califor-
nia [material identified by C.S. Henry].
Chrysoperla downesi (Smith) stat. rev.
(Figs 14, 16)
Chrysopa downesi Smith, 1932: 594. Holotype 9
CANADA: British Colombia, Kelowna
(CNC) [not examined]. [Synonymized with
carnea Stephens by Garland, 1985a: 93.] Stat,
rev.
Chrysopa mohave Banks, 1938: 120. Holotype,
USA (MCZ)[not available for examination].
[Synonymized with C. carnea by Tauber &
Tauber, 1973a: 1164.] Syn. n.
Fore wing (Fig. 14) d\ 11.5 mm; $,
12-12.5 mm. Head marked with black stripe on
gena; red lateral stripe on frons and clypeus; red
post-ocular spot often present. Maxillary palps
black. Antenna shorter than fore wing. Prono-
tum marked with white longitudinal stripe, lat-
eral margins sometimes marked reddish brown;
prothoracic setae short, black. Claw (Fig. 16)
with basal dilation usually less than one-third
length of claw hook (3.7-5.17: 1; mean 4.26) but
specimens with black gradates have a large basal
dilation (2.4-2.7:1). Fore wing narrow (length:
breadth = 3.0-3.5: 1): pointed apically; venation
mostly green but c,lc 2 crossvein black, gradates
green or black; costal setae short (0.4-0.8 mm),
inclined towards wing apex: cell im short, not
reaching 1st Rs crossvein or sometimes meeting
apex of cell: 1st Rs crossvein meets Rs obliquely
or at 90°; 4-6 inner gradates, 5-7 outer gradates.
Genitalia, c/ and 9 and genitalia indistin-
guishable from C. carnea.
Remarks C. downesi and C. plorabunda are
very similar morphologically and it is not always
possible to distinguish the two species. However,
some specimens of downesi, especially those
from the western USA, are marked with a red
post-ocular spot between the eye and vertex.
This marking is never present in plorabunda.
Other characters which serve to distinguish the
two species are the white longitudinal protho-
racic band in downesi. which is yellow in
plorabunda, the basal dilation of the tarsal claw,
which is generally smaller in downesi than
plorabunda, and the width of the fore wing which
tends to be narrower in downesi than
plorabunda. However, these characters are
prone to variation and cannot be relied upon to
separate the species. Earlier workers (e.g.
Tauber & Tauber. 1981) have considered C.
downesi to have a dark green ground colour but,
following courtship song analysis, Henry (1992)
has shown that some populations of C. downesi
include yellow individuals.
Tauber & Tauber (1977a) and Henry (1985a;
1985b) recognised downesi as a valid species on
the basis of ecological evidence. C. downesi
occurs in pine woodland and has a univoltine life
cycle with a summer diapause. In life, the species
has a year-round dark-green ground colour with
a whitish median longitudinal band and is often
covered with an oil-like secretion that is detect-
able in museum specimens. C. plorabunda occurs
in deciduous woodland and open meadow habi-
tats and has a multi-voltine life cycle with no
summer dipause. The species is pale green in the
summer but during the winter, C. plorabunda
becomes reddish brown. Tauber & Tauber
(1977a; 1977b) have postulated that downesi
evolved from plorabunda through a process of
habitat divergence and seasonal isolation. Henry
(1985b) showed that the courtship songs of
downesi and plorabunda were sufficiently differ-
ent to cause reproductive isolation between the
two taxa in the field. However, Henry (1985b)
and Tauber & Tauber (1982a) were able to
152
S.J. BROOKS
hybridise downesi and plorabunda in the labora-
tory, although acceptance rates of heterospecifics
were at low levels (10-30%). Hybridisation does
not appear to occur in the wild (Henry, 1992).
Henry (1985a) reports on the occurrence of
three distinct song morphs of downesi in the
mountains of western USA. Morph Dl is wide-
spread from eastern Canada to western Montana
and south to California. However, this morph
occurs sympatrically with a morphologically
indistinguishable song morph (D2) in the north-
ern Rocky Mountains. D2 co-occurs with a third
song morph (D3) in central California which also
cannot be separated using morphological charac-
ters.
Garland (1985b) states that the female speci-
men bearing the holotype label has a data label
indicating that it was collected in Vernon rather
than in Kelowna as stated in the original descrip-
tion (Smith, 1932).
The true status of C. mohave has been much
discussed in the literature (Tauber & Tauber,
1973a; Henry, 1993). Morphologically the taxon
differs from C. plorabunda in having the cross-
veins black at each end and the gradates com-
pletely black. Tauber & Tauber (1973b; 1982b;
1986; 1987) recognised mohave as a distinct
'strain' of C. plorabunda, with which it hybri-
dises, on ecophysiological and life history
grounds. Henry (1992; 1993) investigated the
courtship song of specimens referrable to C.
mohave and found that they fell into two distinct
groups. One group was indistinguishable from C.
johnsoni, the other was the same as C. downesi.
From a taxonomic point of view it is now impos-
sible to decide whether C. mohave should be
considered synonymous with C. johnsoni or C.
downesi. In order to preserve nomenclatural
stability I have decided to consider C. mohave as
a synonym of C. downesi.
Distribution. Nearctic.
Material examined (31d\ 18$) USA: Cali-
fornia, Colorado, New York, Oregon, Texas,
Utah, Vermont, Wisconsin.
Chrysoperla zastrowi (Esben-Petersen); Holzel,
1989b: 167.
Fore wing (Fig. 24) c? 10.5-13 mm; $
9.5-13 mm. Head sometimes unmarked or
marked with dark brown (occasionally reddish)
spot on gena; reddish brown lateral stripe on
frons and clypeus; maxillary palps brown. Anten-
nae shorter than fore wing. Pronotum with
median longitudinal yellow band; sometimes
with red or brown blotches in anterior corner.
Basal dilation of claw small (4.0-5.0: 1; mean
4.38) Wing venation green; fore wing broad
(length: breadth = 2.8-3.1: 1); costal setae mod-
erately long (0.6-1.2 mm); 1st Rs crossvein
leaves Rs at oblique angle.
Genitaliao" (Figs 26-28). Callus cercus
rounded; 27-37 trichobothria. Gonosaccus with
minute spicules; few gonosetae, about 5-6
arranged in lateral clump; apical membrane of
arcessus short, blunt abruptly curved ventrad.
Genitalia? (Figs 29-31). Callus cercus round;
32 trichobothria. Subgenitale with short basal
extension. Spermatheca wih deep ventral impres-
sion; vela long; duct long.
REMARKS. C. zastrowi is very similar to C. cor-
nea but may be distinguished by the basal dila-
tion of the claw, which is smaller in C. zastrowi
than C. carnea. Tjeder (1966) notes that in C.
zastrowi the 1st Rs crossvein is obliquely angled
whereas in carnea this crossvein is at right-angles
to Rs. However, this character is not always
reliable since several Palaearctic specimens oth-
erwise referrable to carnea have an oblique cross-
vein. Nevertheless, in all southern African
specimens examined the 1st Rs crossvein was
oblique and never at right-angles to Rs. The male
genitalia of C. zastrowi and C. carnea are indis-
tinguishable.
Distribution. Afrotropical.
Material examined (17d\ 13$) . Ascension
I., Namibia, Somalia, St Helena, South Africa,
Sudan, Zambia.
Chrysoperla zastrowi (Esben-Petersen)
(Figs 24-31)
Chrysopa zastrowi Esben-Petersen, 1928a
220.Holotype $ [destroyed], NAMIBIA
Grootfontein (ZMH). Neotype $ NAMIBIA
Orupembe (LUND) [designated by Tjeder
1966: 403.] [not examined].
Chrysopa (Chrysoperla) zastrowi (Esben
Petersen); Tjeder, 1966: 402.
Chrysoperla furcif era (Okamoto)
(Figs 32-39)
Chrysopa furcifera Okamoto, 1914: 61. Lecto-
type $, TAIWAN: Shirin, 31.vii.1907 (Mat-
sumura) (HU) [examined]. [Lectotype
designated by Kuwayama, 1966: 136.]
Chrysopa savioi Navas, 1933c: 4. Holotype $,
CHINA: Kowloon (IZAS) [not examined].
Syn. n.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
153
Chrysopa kulingensis Navas, 1936b: 56. LECTO-
TYPE 0" HERE DESIGNATED. CHINA:
Ruling (IZAS) [examined]. [Synonymized
with savioi by Yang et al., 1990a: 82.]. Syn. n.
Chrysopa (Chrysoperla) furcifera Okamoto;
Tjeder, 1966: 391.
Chrvsoperla furcifera (Okamoto); Tsukaguchi,
1985: 504.
Anisochrysa (Chrysoperla) furcifera (Okamoto);
Holzel, 1970:51.
Chrysoperla savioi (Navas); Brooks & Barnard,
1990:271.
Fore wing (Fig. 32) cf, 11.5-12.5 mm; $,
15 mm. Head with broad black lateral stripe
across gena, clypeus, labrum; in some specimens
clypeus entirely black; large, black or reddish
brown X-marking between antennae with arms
extending onto lateral margins of vertex and
epistomal suture; scape marked with black stripe
on inner and outer lateral surface; maxillary
palps black. Antennae shorter than fore wing;
flagellar segments broad (length: breadth = 2:
1). Pronotum with broad brown lateral band and
yellow longitudinal median band; prothoracic
setae short, black. Claw (Fig. 33) undilated
basally. Fore wing narrow (length: breadth =
3.4-3.7: 1), pointed at apex; venation entirely
green; costal setae short or long (0.6-1.6 mm),
erect or inclined towards wing apex; cell im
relatively broad, 1st Rs crossvein meets im at
apex or subapically; 4-10 inner gradates, 9-10
outer gradates; basal inner gradates not meeting
Psm. Abdominal setae coarse, dense.
Genitalias (Figs 34-36). Dorsal apodeme of
tergite 9 long, extending well beyond callus cer-
cus; callus cercus broadly ovate or circular; 24-36
trichobothria. Lip of sternite 8+9 broad,
rounded. Spinellae absent. Gonosaccus with
minute spicules; few gonosetae, 6-15 in central
clump; arcessus striated dorsally; apex tapering
to point, abruptly curved ventrally. Entoproces-
sus large, crescentic. Gonarcus with large boot-
shaped lateral plate; arch broad. Acumen of
tignum long, narrow, slightly swollen apically.
Genitalia? (Figs 37-39). Callus cercus round;
36 trichobothria. Subgenitale straight basally.
Spermatheca with shallow ventral impression;
duct long; vela long.
Remarks. C. furcifera is one of the most distinc-
tive species in the genus and can be readily
recognized by the extensive black head markings
and narrow, pointed fore wings. Other distin-
guishing characters include the lack of a basal
dilation of the claws, broad flagellar segments.
dense abdominal setae and the inner basal gra-
date not meeting the pseudomedian vein. The
lack of spinellae, striated arcessus, large entopro-
cessus and relatively few gonosetae indicate that
furcifera is related to the carnea-group of species.
Yang & Yang (1992) distinguished the Chinese
species savioi from the Japanese furcifera on the
basis that the clypeus is marked black in savioi
but unmarked in furcifera. However, in one
Chinese example (of nine specimens examined in
this study) the clypeus was only faintly brown
(possibly an immature specimen) and in one
Japanese specimen the clypeus was entirely red-
dish brown. No differences were detected in the
genitalia of these two populations. In view of this
the differences in markings noted by Yang &
Yang (1992) do not seem to me to be sufficient to
regard savioi and furcifera as distinct species.
Geographic variation Specimens examined
from China and Hong Kong differed from other
material in having darker, more extensive head
markings. In these specimens the head markings
are usually black rather than reddish brown. In
addition, the inner stripe on the scape is consid-
erably broader than the outer stripe and the
clypeus is usually (but not invariably) entirely
black. In specimens examined from other regions
the clypeus is unmarked or suffused with red.
Similarly, the Chinese specimens had dark brown
antennae whereas material from other regions
had the antennae pale brown. In addition, it was
noted that the fringe of costal setae was generally
longer in specimens from China and the Philip-
pines (1.2-1.6 mm) than in specimens from
Hong Kong and Japan (0.6-1.0 mm).
Distribution. Bonin I. (Adams, 1959), China,
Hong Kong, Japan, Philippines, Taiwan.
Material examined (6cf , 10$) . China, Hong
Kong, Japan, Philippines, Taiwan.
Chrysoperla iranica (Holzel)
(Figs 40-49)
Chrysopa (Chrysoperla) iranica Holzel, 1967: 33.
Holotype cf, IRAN: Vanak, 1600m, 15km
north of Teheran, l-10.vii.1962 (Vartian)
(HOLZEL COLL) [examined].
Anisochrysa (Chrysoperla) iranica (Holzel); Hol-
zel, 1970:51.
Chrysoperla iranica (Holzel); Brooks & Barnard,
1990: 271.
Fore wing (Fig. 40) d\ 9.5-10 mm; $, 10.5 mm.
Head marked with narrow, dark brown stripe on
gena and lateral clypeus. Maxillary palps black
154
S.J. BROOKS
dorsally. Antennae shorter than fore wing.
Pronotum marked with median longitudinal
white/yellow stripe; prothoracic setae short, pale.
Claw (Fig. 41) with small basal dilation, about
one-third length of claw hook (2.72: 1). Fore
wing broad (length: breadth = 2.7-3.1: 1),
rounded apically; costal setae quite long
(0.8-1.0 mm), inclined towards wing apex; 4-6
inner gradates, 5-7 outer gradates; gradates
green; crossveins black at each end, except costal
and Psm crossveins which are green at the wing
margins; anal veins green; cell im variable in
length. Abdominal tergites with white/yellow
median longitudinal band.
Genitalias (Figs 42-46). Lip of sternite 8+9
elongate, truncate apically with subapical con-
striction. Apodeme of tergite 9 long, extends far
beyond callus cercus; callus cercus narrow,
ovate; 27 trichobothria. Spinellae absent.
Gonosetae few, 5-9 in lateral clump. Arcessus
with dorsal striations; apex blunt; apical mem-
brane short, bent ventrad at 90°. Entoprocessus
large, crescentic. Gonarcus with lateral plate
broad, reniform; arch broad. Acumen of tignum
narrow, rounded apically.
Genitalia? (Figs 47-49). Callus cercus ovate;
30 trichobothria. Subgenitale with narrow, short
basal extension. Spermatheca with shallow ven-
tral impression; duct long; vela long.
Remarks. C. iranica belongs to the carnea-
group of species. However, iranica can be distin-
guished from these species by the broad fore
wing with rounded apex and, in males, by the
extraordinary elongation of the lip on sternite
8+9.
Distribution. Iran.
Material examined (3cf , 3$) Iran.
Chrysoperla mediterranea (Hdlzel)
(Figs 50-58)
Anisochrysa (Chrysoperla) mediterranea H61-
zel, 1972: 81. Holotype cf, TUNISIA: Biserta,
100m, 9.viii.l971 (Gepp) (HOLZEL COLL.)
[not examined].
Anisochrysa mediterranea (Holzel); Monser-
rat, 1977: 130.
Chrysoperla mediterranea (Holzel); Aspock et
al., 1980: 269.
Fore wing (Fig. 50) cf , 8.5-9.5 mm; $, 11 mm.
Head sometimes entirely suffused red or marked
with red/brown stripe on gena, frons and clypeus;
labrum sometimes red; red post-ocular spot
sometimes present. Maxillary palps black dor-
sally or unmarked. Antennae shorter than fore
wing. Pronotum marked with longitudinal
median yellow stripe and sometimes with brown
median dorso-lateral spot; prothoracic setae
short, black or pale. Claw (Fig. 51) with minute
basal dilation (8.25: 1). Fore wing narrow
(length: breadth = 3.2^4.0: 1); venation green;
costal setae short (0.4-0.6 mm), inclined towards
wing apex; cell im short, apex of cell not reaching
1st Rs crossvein; 4-6 inner gradates, 5-7 outer
gradates.
Genitaliao" (Figs 52-55). Membranous lip of
sternite 8+9 broad, rounded. Apodeme of terg-
ite 9 long, extending well beyond callus cercus;
23-24 trichobothria. Spinellae absent. Gonosetae
few, 5-6 in lateral clump. Arcessus with dorsal
striations; straight but extreme apex bent ventrad
at 90°, apex blunt. Entoprocessus large, crescen-
tic. Lateral plate of gonarcus large, reniform;
arch broad. Acumen of tignum long, rounded
(not swollen) apically.
Genitalia? (Figs 56-58). Callus cercus ovate;
28 trichobothria. Subgenitale with short basal
projection. Spermatheca narrow; ventral impres-
sion deep; duct long.
Remarks. C. mediterranea is apparently closely
related to carnea and shares several characters
with that species. Like carnea the gena is marked
with a brown stripe, the wing venation is green,
costal setae short and intramedian cell short.
Similarly, the male genitalia of both species have
many characters in common: spinellae are
absent, the arcessus is dorsally striated and there
are few gonosetae. However, the wings are nar-
rower in mediterranea and the basal dilation of
the claw much smaller than in carnea and other
related species. On the basis of the these charac-
ters it seems justified to continue to treat mediter-
ranea as a distinct species.
Duelli (1987) described a population of green
lacewings referrable to mediterranea from Swit-
zerland. The population occurred sympatrically
with carnea but was ecologically separated since
carnea was confined to oak whereas mediterranea
was found on pine. In the laboratory the taxa
interbred and produced fertile hybrids.
Cianchi & Bullini (1992) report that their
electrophoretic work on this Swiss population of
mediterranea suggests it is genetically distinct
from other populations of mediterranea
(although they do not state which populations
they examined) and should be considered a dis-
tinct taxon. However, until comparable data is
available from a wide range of populations of this
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
155
species it is premature to formally recognise the
taxa as distinct species.
Distribution. North-west Africa, south-west
Europe, Hungary (Szentkiralyi, pers comm.),
Switzerland (Duelli, 1987).
Material examined (2cf, 1?) Algeria, main-
land Spain, Majorca.
Chrysoperla nipponensis (Okamoto) stat.
rev.
(Figs 59-67)
Chrysopa nipponensis Okamoto, 1914: 65.
Lectotype a", JAPAN: Kyushyu, Kumamoto,
4.iv.l907 (Kawamura) [Lectotype designation by
Kuwayama, 1966: 135. ](HU) [examined].
Chrysopa kurisakiana Okamoto, 1914: 71.
Lectotype $, JAPAN: Mt Fuji, 23. vi. 1911 [Lec-
totype designation by Kuwayama, 1966: 135.]
(HU) [not examined]. [Synonymized by
Kuwayama, 1962: 1 1.]
Chrysopa ilota Banks, 1915: 629. Holotype $,
PHILIPPINE IS: Luzon, Mt Makiling {Baker)
(MCZ) [examined]. Syn. n.
Chrysopa kolthoffiNavds, 1927b: 3. Lectotype
$, CHINA: Kiangsu (Kolthoff) [Lectotype desig-
nation by Leraut, 1991: 79.] (MNHN) [exam-
ined]. [Synonymized with carnea by Tjeder,
1936: 32; raised to valid species by Leraut, 1992:
63.] Syn. n.
Chrysopa kolthoff i var. adaptata Navas, 1934a:
2. Holotype $, CHINA: Jiangsu Prov., Hufu
(Loubou), 3.viii.l933 (IZAS) [not examined].
Syn. n.
Chrysopa sinica Tjeder, 1936: 29. Holotype
Cf, CHINA: South Kansu, Ardjuan, 2900m,
8.ix.l930 (Sjostedt) (NRS) [examined]. Syn. n.
Chrysopa (Chrysoperla) sinica Tjeder; Tjeder,
1966: 391.
Anisochrysa (Chrysoperla) sinica (Tjeder);
Holzel, 1970: 51.
Chrysoperla nipponensis (Okamoto); Tsuk-
aguchi, 1985: 504 [as synonym of Chrysoperla
carnea (Stephens)].
Anisochrysa (Chrysoperla) sinica (Tjeder);
Holzel, 1970: 51.
Chrysopa (Chrysoperla) sinica Tjeder; Tjeder,
1966: 391.
Fore wing (Fig. 59) 0\ 10.5-13 mm; $>,
12.5 mm. Head marked with black/brown stripe
on gena and lateral clypeus; lateral frons some-
times marked with red stripe. Maxillary palps
black dorsally. Antennae shorter than fore wing.
Pronotum marked with median yellow longitudi-
nal band and pale brown lateral stripe; protho-
racic setae long or short, pale. Claw (Figs 60, 61)
with large basal dilation, about half length of
claw hook (2.07-2.42: 1; mean 2.22). Fore wing
relatively broad (length: breadth = 2.9-3.1: 1);
apex pointed; costal setae long (1.0-2.4 mm),
inclined towards wing apex; cell im short, not
meeting 1st Rs crossvein; 1st Rs crossvein usually
at right-angles to Rs; 4-6 inner gradates, 6-7
outer gradates; gradates black; crossveins dark at
each end except costal and Psm crossveins which
are unmarked at the wing margins.
Genitaliao" (Figs 62-64). Dorsal apodeme of
tergite 9 long, extending beyond callus cercus;
callus cercus broad, ovate; 27-33 trichobothria.
Membranous apex of sternite 8 + 9 broad,
rounded. Spinellae absent. Gonosaccus with
minute spicules; gonosetae few, 5-7 in lateral
clump. Arcessus with dorsal striations; apex
blunt, curved ventrally at 90°. Entoprocessus
large, crescentic. Gonarcus lateral plate broadly
reniform; arch broad. Acumen of tignum long,
narrow, rounded apical ly.
Genitalia9 (Figs 65-67). Callus cercus round;
32 trichobothria. Subgenitale with short, narrow
basal extension. Spermatheca with shallow ven-
tral impression; duct long; vela long.
Remarks. C. nipponensis is closely related to
carnea and the male genitalia of the two species
appear to be indistinguishable. However, there
are consistent differences in the external mor-
phology that suggest they should be treated as
distinct species. In nipponensis the gradates are
black, the costal setae relatively long and the
basal dilation of the claw is about half the length
of the claw hook. In addition, the gena and
lateral clypeus is heavily marked dark brown/
black, the prothoracic setae are pale and the
acumen of the tignum is relatively long (Fig. 63).
In carnea the gradates are green, costal setae
generally short and the basal dilation of the claw
is about one-third the length of the claw hook. In
most examples of carnea examined the genal
markings are less extensive and paler, the pro-
thoracic setae are black and the acumen is
shorter.
Yang & Yang (1990a) examined the male type
of Chrysopa chusanina Navas and commented
that the genitalia indicated the species should be
transferred to Chrysoperla. They also considered
that the species may be a synonym of C. sinica
Tjeder. Unfortunately, I have not been able to
borrow the type of C. chusanina so cannot con-
firm this view. However, the original description
of C. chusanina (Navas, 1933c) does not
resemble that of sinica Tjeder. Navas describes
156
S.J. BROOKS
the palps as yellow, a red arcuate stripe in front
of the antennae, a red lateral stripe on the head
and a red lateral stripe on the pronotum. None of
these markings occur in sinica. However, without
access to the type material it is impossible to
decide the true affinities of the species.
Thierry et al., (1992) and Leraut (1992) incor-
rectly refer to a French morph of carnea as
kolthoffi.
Geographical variation. Korean specimens
and some of the Chinese material examined have
a red stripe on the lateral frons that is absent in
Japanese material.
The Mongolian paratypes of sinica have the
gena marked with a narrow, pale brown stripe
and in one specimen the gradate crossveins were
green, not marked black.
Distribution. China, Japan, Korea, Mongolia,
Philippines.
Material examined (9d\ 15$) China, Japan,
Korea, Mongolia.
Chrysoperla renoni (Lacroix)
(Figs 68-75)
Chrysopa renoni Lacroix, 1933: 149. Lecto-
typecf, FRANCE: Deux-Sevres, Amure,
29.vi.1930 {Lacroix) [Leraut, 1991: 72. Lecto-
type designation.] (MNHN) [examined].
Chrysoperla ankylopteryformis Monserrat &
Diaz-Aranda,1989: 261. Holotype 0", SPAIN:
Almeria, Rambla de Morales (Cabo de Gata),
10m, 12.viii.1988 [MONSERRAT COLL]
[examined]. Syn. n.
Fore wing (Fig. 68) d\ 10.5 mm; $, 11.5 mm.
Head marked with brown stripe on gena and
lateral clypeus. Maxillary palps unmarked or
black dorsally. Antenna very short, little more
than half length of fore wing. Pronotum marked
with median longitudinal white/yellow band,
vague broad pale reddish medio-lateral red
stripe, narrow antero-ventro-lateral black stripe;
prothoracic setae very long, pale; setal follicles
sometimes marked black. Claws with minute
basal dilation (10.25: 1), claw tooth elongate
(Fig. 69). Fore wing variable in width (length:
breadth = 2.8-3.7: 1); rounded apically; vena-
tion mainly green with costal crossveins some-
times black at subcosta; costal setae long
(1.8-2.4 mm), erect; cell im short, not reaching
1st Rs crossvein; 1st Rs crossvein meets Rs
obliquely; 4-6 inner gradates, 5-6 outer gra-
dates.
Genitaliao" (Figs 70-72). Dorsal apodeme of
tergite 9 long, extending well beyond callus cer-
cus; callus cercus broad, ovate; 30 trichobothria.
Lip of sternite 8+9 narrow, prominent. Spinellae
absent. Gonosetae few, 5 arranged in lateral
clump. Arcessus broad with dorsal striations;
apex blunt, curved at 90°. Entoprocessus large,
crescentic. Gonarcus with broad lateral plate;
arch broad. Acumen of tignum broad, not swol-
len apically.
Genitalia? (Figs 73-75). Callus cercus broad,
ovate; 26 trichobothria. Subgenitale with short,
broad basal extension. Spermatheca with shallow
ventral impression; duct long; vela short.
Remarks. On the basis of the male genitalia, C.
renoni belongs to the carnea-group of species. I
could detect no useful characters in the gonarcus-
complex to distinguish renoni from other species
in this group. However, the acumen of the
tignum is shorter and broader in renoni than in
males of carnea. In females of renoni the basal
lobe of the subgenitale is longer than usual in
other members of the carnea-group. From other
Chrysoperla species renoni can be readily identi-
fied by the long prothoracic and costal setae and
the minute basal dilation of the claw. The claws
of C. mediterranea also have a minute basal
dilation but the claw hook is much shorter than in
renoni and not at right angles to the claw base. In
addition, the fore wings of mediterranea are
much narrower than in renoni and are pointed
apically rather than rounded.
The present synonymy between C. renoni and
C. ankylopteryformis confirms the suggestions of
Leraut (1991) and Aspock (1992).
Distribution. France, Spain.
Material examined (3d\ 3$, 1 without abdo-
men)
France (including lectotype of renoni), Spain
(holotype and two paratypes of ankylopteryfor-
mis).
Chrysoperla oblita (Holzel)
(Figs 76-83)
Anisochrysa (Chrysoperla) oblita Holzel,
1973:342. Holotype C\ NEPAL: Jiri,
12. vi. 1964 (Remane) (ZSBZ) [examined].
Chrysoperla oblita (Holzel); Brooks & Barnard,
1990: 271.
Fore wing (Fig. 76) cf, 10.5-11 mm; $,
10.5-13 mm. Head marked with broad reddish
band across gena, lateral clypeus and labrum,
markings sometimes extending onto lateral frons
and medio-lateral area of clypeus; red post-
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
157
ocular spot. Maxillary palps unmarked. Anten-
nae as long as fore wing. Pronotum with median
longitudinal yellow band and red spot in anterior
corner; prothoracic setae long or short, pale.
Claw (Fig. 77) with large basal dilation, about
half length of claw hook (1.88-2.14: 1; mean
2.01). Fore wing with gradates black; other cross-
veins dark each end, except Psm crossveins
entirely green; upper branch of forking marginal
veins black; anal veins green; apex of wing
rounded; costal setae long (1.0-1.2 mm), erector
slightly inclined towards wing apex; 5-7 inner
gradates, 6-8 outer gradates; cell im long, 1st Rs
crossvein meets im at apex or subapically; 1st Rs
crossvein leaves Rs at right-angles or obliquely.
Genitalias (Figs 78-80). Lip of sternite 8+9
broad, truncate, at 45° to sternite. Apodeme of
tergite 9 long, extending beyond callus cercus;
callus cercus broad, ovate; 28-30 trichobothria.
Spinellae absent. Gonosaccus with minute spi-
cules; gonosetae few, 9-12 in lateral clump.
Arcessus with dorsal striations; apex blunt,
curved evenly ventrad. Entoprocessus large,
crescentic. Gonarcus broad, lateral plate foot-
shaped; arch broad. Acumen of tignum short,
very broad, rounded apically.
Genitalia? (Figs 81-83). Callus cercus
rounded; 25 trichobothria. Subgenitale concave
basally with median projection. Spermatheca
with shallow ventral impression; duct long; vela
short.
Remarks. C. oblita belongs to the group of
species related to carnea that have a striated
arcessus and lack spinellae in the male genitalia.
However, the species differs from others in the
group by having relatively more gonosetae in the
male genitalia {carnea has only 5-6 in each lateral
group), the gena marked red instead of brown
and a large basal dilation of the claw. C. oblita is
readily identifiable by the black gradate cross-
veins, long costal setae and the broad acumen in
the male genitalia.
The holotype of oblita is now in poor condi-
tion. Both fore wings are broken off at the base.
The apex of the abdomen is preserved in a vial of
glycerine but the internal genitalic components
(gonarcus, arcessus and tignum) are all missing.
However, Holzel's (1973) figures and desciption
of the type are sufficient for positive identifica-
tion of the species.
Distribution. Highlands of northern Thailand,
Nepal and north-west India. All specimens were
collected between 1380 and 1700m.
Material examined (4q\ 8$) . Thailand:
Chiang Mai Province. India: Assam, Khasia
Hills. Nepal.
Chrysoperla annae sp. n.
(Figs 84-89)
Fore wing (Fig. 84) q\ 13.5 mm; 9 unknown.
Head marked with reddish brown stripe on gena
and lateral clypeus, red stripe on frontal suture.
Maxillary palp black dorsally. Antenna shorter
than fore wing. Pronotum with yellow longitudi-
nal median band; setae short, black. Claw (Fig.
85) with small basal dilation, one-quarter or less
than claw hook length (3.25-9.75: 1; mean 6.5).
Fore wing narrow (length: breadth = 3.1-3.3: 1),
venation green; costal setae moderately long
(0.8-1.0 mm), inclined towards wing apex; 1st Rs
crossvein meets im subapically; 6-7 inner gra-
dates, 7-9 outer gradates.
Genitalia^ (Figs 86-89). Dorsal apodeme of
tergite 9 short; callus cercus broadly ovate; 25-28
trichobothria. Lip of sternite 8+9 broad, trun-
cate when viewed ventrally. Spinellae absent.
Gonosaccus with 12-20 gonosetae in lateral
group. Arcessus narrow, straight, striated dor-
sally; apex rounded, straight or curved ventrad at
90°. Entoprocessus small, narrow, crescentic.
Lateral plates of gonarcus narrow, reniform;
arch broad. Acumen of tignum moderately long,
narrow not swollen apically.
Remarks Chrysoperla annae belongs to the
carnea-group of species. However, the new spe-
cies possesses several distinctive characters that
distinguish it from other species in the group. In
the male genitalia the arcessus, entoprocessus
and the lateral plates of the gonarcus are nar-
rower than in carnea and there are 12-20 gonose-
tae, instead of six, in each lateral group. The
relatively narrow fore wing with rounded apex is
also characteristic of the species.
Distribution. Burma, Tibet.
Material examined (2cf ) Holotype d\ Tibet:
Tsaidam (Prejevdoky) (BMNH). Paratype d\
Burma: Tongkruk, 9,000, 20.vii.1931 (Kingdon
Ward) (BMNH).
Chrysoperla gallagheri Holzel
(Figs 90-98)
Chrysoperla gallagheri Holzel, 1989a: 60. Holo-
type d\ OMAN: Jabal Shams, 2500m, 23.15N/
57.15E, 16.vii.1987 {Gallagher) (HOLZEL
COLL.) [examined].
158
S.J. BROOKS
Fore wing (Fig. 90) C\ 9.5-10.5 mm; ?, 12 mm.
Head marked with broad red stripe across gena,
lateral frons, lateral clypeus; red post-ocular
stripe; head sometimes suffused entirely red;
lateral scape and pedicel with red stripe. Maxil-
lary palp unmarked. Antenna about same length
as fore wing. Pronotum unmarked, pale median
longitudinal stripe absent but yellow/green
median stripe present on pteronotum; protho-
racic setae long, black; follicles unmarked. Claw
(Fig. 91) with small basal dilation, less than a
quarter length of claw hook (5.5: 1). Abdomen
marked with dorsal median yellow stripe on
tergites 1-3. Fore wing (length: breadth =
2.9-3.0: 1), pointed apically; costal setae rela-
tively long (1.0 mm), inclined towards wing
apex; 4—5 inner gradates, 5-7 outer gradates;
gradates black; crossveins black each end; 1st Rs
crossvein meets cell im at apex; 1st Rs crossvein
straight or oblique.
GenitaliacT (Figs 92-95). Lip of sternite 8+9
narrow, rounded apically; apex of sternite 8+9
narrowed to form prominent tubercle. Apodeme
of tergite 9 long, extending far beyond callus
cercus; callus cercus moderately narrow, ovate;
26 trichobothria. Spinellae absent. Gonosetae
numerous, formed into lateral group. Arcessus
with dorsal striations; apical membrane long,
narrow, curved at 45°, rounded at tip. Entopro-
cessus short, narrow, crescentic. Acumen of
tignum short, broad, swollen; arms of tignum
narrow.
Genitalia9 (Figs 96-98). Callus cercus ovate;
33 trichobothria. Subgenitale with short, narrow
basal extension. Spermatheca with deep ventral
impression; duct long; vela short.
Remarks. Like C. pudica and mutata, the gra-
dates of gallagheri are black. However, gallagh-
eri may be distinguished from these species by
the small basal dilation of the claw, the red stripe
on the scape and, in males, by the absence of
spinellae.
Distribution. Oman.
Material examined (2d\ 2$) Oman.
Chrysoperla comans (Tjeder)
(Figs 99-108)
Chrysopa (Chrysoperla) comans Tjeder, 1966:
408. Holotype Q\ ZIMBABWE: Umtali,
14. i. 1945 (Wright)(TM) [not examined].
Chrysoperla comans (Tjeder); Holzel, 1989b:
175.
Chryoperla manselli Holzel, 1989b: 176. Holo-
type cf , SOUTH AFRICA: Transvaal, Vrede-
huis gardens of Union Buildings, 25.45S/
28.12E, l.x.1979, at light (Mansell) (NCI)
[examined]. Syn. n.
Fore wing (Fig. 99) cf, 10.5-11.5 mm; $,
12.0-13.5 mm. Head usually marked with red
lateral stripe on gena and clypeus; red post-
ocular stripe. Maxillary palps unmarked. Anten-
nae about as long as fore wing. Pronotum
marked with red spot in anterior corner and
yellow median, longitudinal band; prothoracic
setae long (2.2-3.0 mm), pale. Claw (Fig. 100)
with large basal dilation, about two-thirds length
of claw hook (1.59-1.71: 1; mean 1.65). Fore
wing broad, with brown gradates; costal setae
long (2.2-3.0 mm), erect; 5-8 inner gradates, 6-9
outer gradates; 1st Rs crossvein meets im apically
or sub-apically.
Genitaliao" (Figs 101-105). Sternite 8+9 elon-
gate; lip small, broad. Callus cercus small, nar-
row, ovate; 26 trichobothria. Apodeme of tergite
long, often branched, extending considerably
beyond callus cercus. Spinellae absent. Gonose-
tae short, numerous, evenly dispersed across
gonosaccus. Arcessus narrow, apical membrane
long, blunt, slightly curved or recurved ventrad,
not striated dorsally. Entoprocessus small cres-
centic. Lateral plate of gonarcus ovate or boot-
shaped. Acumen of tignum short, grossly
swollen.
Genitalia9 (Figs 106-108). Callus cercus
ovate; 26 trichobothria. Subgenitale with short
basal projection. Spermatheca with deep ventral
impression; duct long; vela long.
Remarks. C. comans belongs to the group of
Afrotropical broad-winged species with long,
erect costal setae. From nyerina and plicata it can
be distinguished by the absence of spinellae. C.
comans can be distinguished from the Madagas-
can species, decaryana, by the narrow arcessus,
short, swollen acumen and the narrow, elongate
apex of sternite 8+9. In decaryana the arcessus is
broad, the acumen narrow and sternite 8+9 is
not elongated apically.
When he described manselli, Holzel (1989b)
remarked on the similarities between manselli
and comans, in particular the elongation of stern-
ite 8+9, which does not occur elsewhere in the
genus. However, he then went on to say that the
gonarcus complex and tignum were totally differ-
ent in the two taxa. Nevertheless, on
re-examination of the types of manselli I can see
no significant differences in the genitalia from
that described by Tjeder (1966) for comans.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
159
Indeed Holzel's (1989b) figure of the acumen of
manselli shows it less swollen than it is in fact.
Distribution. Kenya, Zimbabwe, South
Africa.
Material examined (3cf, 39) ■ South Africa,
Kenya.
Chrysoperla decaryana (Navas)
(Figs 109-114)
Chrysopa decaryana Navas, 1934b: 60. Holo-
typecf, MADAGASCAR: Tananarive, 1921
(Decary) (MNHN) [examined].
Chrysoperla decaryana (Navas); Holzel. 1989b:
169.
Fore wing (Fig. 109) d\ 13 mm; 9 ur| known.
Head marked with broad red lateral stripe on
gena, frons and clypeus; red post-ocular stripe.
Maxillary palp with black dorsal stripe. Antenna
shorter than fore wing. Pronotum marked with
yellow/white median longitudinal band and red
medio-lateral spot; prothoracic setae long, pale.
Basal dilation of claw large, about half length of
claw hook (2.25: 1) (Fig. 110). Fore wing broad
(length: breadth = 3: 1); costal setae long
(1.6 mm), slightly inclined apically; Rs crossveins
sinuate; gradates black and other crossveins dark
at each end except costals at costa and Psc
crossveins entirely green; 1st Rs crossvein meets
im at apex.
Genitalias (Figs 111-114). Lip of sternite 8+9
narrow, tapering apically, prominent; callus cer-
cus narrow, ovate; 32 trichobothria; apodeme of
tergite 9 short, not extending beyond callus cer-
cus, branched. Spinellae absent. Gonosaccus
with minute spicules. Gonosetae few (about 14)
in median group. Entoprocessus minute, crescen-
tic. Arcessus lacking dorsal striations; short,
broad, straight; apex blunt, rounded. Gonarcus
with broad arch; basal dilation narrow wedge.
Acumen of tignum long, parallel sided, rounded
apically; arms narrow.
Remarks. C. decaryana belongs to the group of
Afrotropical species with broad wings and long
costal setae. It is probably most closely related to
comans Tjeder since both species lack spinellae
in the male genitalia. However, from this species
decaryana can be distinguished by the broad
arcessus and relatively narrow acumen.
Chrysoperla siamensis sp. n.
(Figs 115-123)
Fore wing (Fig. 115) cf, 11-12.5 mm; $, 14 mm.
Head sometimes unmarked or marked with red-
dish spot on lateral clypeus, extending to medio-
lateral area; red spot on lateral frons which
sometimes extends posteriorly as stripe between
eye and scape; red post-ocular stripe; gena
unmarked, or sometmes with red spot. Maxillary
palps unmarked. Antennae shorter than fore
wing. Pronotum marked with yellow median
longitudinal band, bordered yellow/orange,
sometimes marked with red medio-lateral spot;
prothoracic setae short, pale. Claw (Fig. 116)
with large basal dilation, over half length of claw
hook (1.86: 1). Fore wing broad (length: breadth
= 2.8-3.0: 1); rounded apically; costal setae long
(1.0-1.4 mm), erect; 1st Rs crossvein leaves Rs at
oblique angle; 1st Rs crossvein meets cell im
subapically; Rs crossveins sinuous; gradates
green or greyish with surrounding membrane
suffused grey, other veins green; 4-7 inner gra-
dates, 6-8 outer gradates.
Genitalias (Figs 117-120). Lip of sternite 8+9
broad, rounded. Dorsal apodeme of tergite 9
extending beyond callus cercus; callus cercus
broad, ovate; 23-38 trichobothria. Spinellae
absent. Gonosetae numerous in median group.
Arcessus lacking dorsal striations; narrow, curv-
ing in apical half to 90°; membranous tip very
long and narrow, blunt apically. Entoprocessus
very small, narrow, crescentic (not visible in
some specimens examined). Gonarcus with lat-
eral plates broad, ovate; arch broad. Acumen of
tignum short, swollen apically.
Genitalia? (Figs 121-123). Callus cercus
ovate; 28 trichobothria. Subgenitale with narrow
basal extension. Spermatheca with shallow ven-
tral impression; duct short; vela short.
Remarks. C. siamensis is unusual amongst
Chrysoperla species in lacking both spinellae and
dorsal striations on the arcessus in the male
genitalia. The only other species that also lack
these characters are comans, decaryana and gal-
lagheri. Other distinctive characters of siamensis
are the broad, rounded fore wing with a grey
suffusion around the gradate crossveins, the rela-
tively long intramedian cell, sinuous Rs cross-
veins and short pale prothoracic setae. In
addition, the arcessus is very narrow and the lip
of sternite 8+9 unusually broad.
Distribution. Thailand.
Material examined (1 cf). Madagascar.
Material examined (5$, 2$) Holotype cf,
160
S.J. BROOKS
Thailand: Chiang Mai Prov., Mae Rim to
Samoeng Road, 24 Km post, 1100m, 29.U989
(Cotton) (BMNH). Paratypes. Thailand: 2cf,
Korat Prov., Khao Yai National Park, Khao
Kieow, 1200m, l.iii.1989 (Cotton); 1$, Chiang
Mai Prov., Doi Suthep-Pui National Park,
1380m, 24.ii. 1989 (Cotton); lcf, 1$, Doi Suthep-
Pui National Park, 1460m, 26.iv-10.v.l989 (Cot-
ton); lcf, Doi Suthep-Pui National Park, 1200m,
10-11. xi. 1988 (Bradley, Lewvanich, Fletcher)
(BMNH).
Chrysoperla nyerina (Navas)
(Figs 124-132)
Chrysopa nyerina Navas, 1933a: 210. Holotype
d\ KENYA: Nyeri, 9.vi.l932 (MNHN)
[examined].
Chrysopa socia Navas, 1936a: 116. Lectotype $,
KENYA: Marakwet, Elgeyo Escarpment,
2500m, 1932-33 (Arambourg, Chappuis &
Jeannel) [Lectotype designation by Holzel,
1989b: 173.] (MNHN) [examined]. Syn. n.
Chrysoperla nyerina (Navas); Holzel, 1989b:
172.
Chrysoperla socia (Navas); Holzel, 1989b: 173.
Fore wing (Fig. 124) cf, 13-14.5 mm; $,
13.5-15 mm. Head occasionally unmarked or
marked with broad red lateral stripe on gena and
clypeus; red post-ocular spot. Maxillary palps
black dorsally, sometimes unmarked. Specimens
examined from Uganda had a red lateral stripe
on the scape, otherwise scape unmarked. Prono-
tum with longitudinal yellow median band,
sometimes with red spot in anterior corner; pro-
thoracic setae very long, pale. Claw (Fig. 125)
with large basal dilation, about two-thirds length
of claw hook (1.77-1.89: 1; mean 1.83). Fore
wing broad (length: breadth = 2.6-2.8: 1); costal
setae long (1.8-2.6 mm), erect; gradates black
and other crossveins dark at each end (except Psc
crossveins which are green at wing margin); 6
inner gradates, 6-7 outer gradates; im long, 1st
Rs crossvein meets im sub-apically. Abdomen
with pair of red dorsal spots on basal segments.
Genitalias (Figs 126-129). Lip of sternite 8+9
broadly rounded. Apodeme of tergite 9 long,
extends far beyond callus cercus; callus cercus
broad, ovate; 21-36 trichobothria. Few (about
20) minute spinellae, each spine on an individual
tubercle. Gonosetae numerous, evenly distrib-
uted on gonosaccus. Entoprocessus small, cres-
centic. Arcessus long, narrow with dorsal
striations; weakly curved ventrad; truncate api-
cally. Gonarcus with narrow lateral plate. Acu-
men of tignum long, parallel sided, rounded
apically.
Genitalia? (Figs 130-132). Callus cercus
ovate; 25 trichobothria. Subgenitale broad. Sper-
matheca narrow; ventral impression deep; vela
short; duct long.
Remarks. C nyerina is probably closely related
to plicata since this is the only other species in the
genus with simple spinellae. However, there are
several characters that will distinguish the spe-
cies. In nyerina the basal dilation of the claw is
larger than in plicata. In the male genitalia the
apex of the acumen is not swollen in nyerina and
it has fewer and smaller spinellae than plicata. In
plicata, unlike nyerina, the spinellae are not set
in tubercles.
Distribution. Kenya, Sudan, Uganda.
Material examined (4cf , 6$) Kenya, Sudan,
Uganda
Chrysoperla plicata (Tjeder)
(Figs 133-141)
Chrysopa (Chrysoperla) plicata Tjeder, 1966:
406. Holotype cf, ZIMBABWE: Marandellas,
Eirene Farm, x.1960 (NHMZ) [examined].
Chrysopa plicata (Tjeder); Holzel, 1989b: 168.
Fore wing (Fig. 133) cf, 9.5-12.5 mm; $,
12.5-13 mm. Head marked with strong red spot;
red post-ocular stripe. Maxillary palps
unmarked. Antennae about as long as fore wing.
Pronotum marked with red stripe in each corner
and red centro-lateral spot; median longitudinal
yellow band; prothoracic setae pale, long or
short. Claw (Fig. 134) with large basal dilation,
about two-thirds length of claw tooth (1.58: 1);
rounded posteriorly. Fore wing marked with
black spot on tympanal organ; crossveins black at
each end (except marginal crossveins at wing
margin); gradates black; 3-7 inner gradates, 4-8
outer gradates; costal setae long (1.0—1.8 mm),
slightly inclined; im variable in length.
Genitalias (Figs 135-138). Apodeme of terg-
ite 9 variable, usually not extending beyond
callus cercus; 21-30 trichobothria; lip of sternite
8 + 9 prominent, strongly tapered apically when
viewed ventrally. Spinellae present, numerous,
single broad spines not mounted on tubercle.
Gonosetae numerous, evenly distributed across
gonosaccus. Arcessus with dorsal striations;
blunt apically, weakly curved. Entoprocessus
short, narrow, triangular. Gonarcus with wide
arch and median projection; lateral plates broad,
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
161
reniform. Acumen of tignum short, broad, bul-
bous apically.
Genitalia? (Figs 139-141). Callus cercus
rounded; 29 trichobothria. Subgenitale with
large basal extension. Spermatheca narrow; ven-
tral impression shallow; vela short; duct long.
Remarks. Males of C. plicata are unusual
amongst Chrysoperla species in that the arcessus
is striated and spinellae are present. The only
other species to share these characters is nyerina.
C. plicata also shares with nyerina simple spinel-
lae in the male genitalia. Unlike nyerina, how-
ever, the spinellae of plicata arise directly from
the gonosaccus and are not raised on tubercles.
In most other species in the genus which possess
spinellae, these structures are composed of sev-
eral spines positioned on each tubercle. C. pli-
cata may be distinguished from other
Chrysoperla species by the strongly defined red
spot on the gena and the three red lateral stripes
on the pronotum. In addition, the claw is
strongly reflexed and has a large basal dilation.
Tjeder's (1966) figure of the male genitalia of
plicata shows the apex of the arcessus strongly
hooked. In the specimens examined during this
study all had the arcessus only weakly curved.
Unfortunately, the genitalia of the holotype have
been slide mounted in such a way that it is
impossible to see the extent of the curvature of
the arcessus in the holotype.
Distribution Afrotropics.
Material examined (9o", 3$) Mozambique,
Sene-Gambia, Somalia, South Africa, Zaire,
Zambia, Zimbabwe.
Chrysoperla insulata (Fraser)
(Figs 142-148)
Chrysopa insulata Fraser, 1957: 26. Syntypesd"
$, REUNION: Hellbourg; forest track in the
forest of Belouve; Bebour & Plaine des Caf-
res, Piton Manuel, I. 1955 (Paulian) (IRSM &
BMNH) [ld\ 1$ syntype in BMNH exam-
ined].
Chrysoperla insulata (Fraser); Brooks & Bar-
nard, 1990: 271.
Fore wing (Fig. 142) o\ 13 mm; $, 16.5 mm.
Head marked with black lateral stripe on
clypeus. Maxillary palps black. Antennae shorter
than fore wing. Pronotum with median longitudi-
nal yellow band; prothoracic setae short, pale.
Claw (Fig. 143) with large basal dilation, over
half the length of claw hook (2.21: 1). Fore wing
broad (length: breadth = 28.-2.9: 1), pointed
apically; costal setae long (2.4-2.8 mm), erect;
im long, apex extends beyond 1st Rs crossvein;
gradates black; 8-9 inner gradates, 9-10 outer
gradates; other crossveins darkened at each end.
Genitalias (Figs 144-145). Lip of sternite 8+9
broad, rounded; apodeme of tergite 9 short, not
extending beyond callus cercus; 31 trichobothria.
Spinellae present, numerous, 2-3 spines on
tubercle. Gonosetae numerous, arranged in lat-
eral group. Arcessus narrow; lacking dorsal stria-
tions; strongly curved ventrad; apical membrane
long, narrow, truncate at apex. Entoprocessus
minute, narrow, straight. Gonarcus narrowly
expanded basally. Tignum absent.
Genitalia? (Figs 146-148). Callus cercus
ovate; 39 trichobothria. Subgenitale broad with
narrow basal extension. Spermatheca with mod-
erate ventral impression; vela short; duct long.
Remarks. C. insulata is unique amongst
Chrysoperla species because the tignum is absent
in the male genitalia. The species is also charac-
terized by the relative lack of head markings,
only the clypeus is marked with a lateral black
stripe, and the broad distinctly pointed fore
wings. C. insulata is the largest Chrysoperla
species with a fore wing length of 16.5 mm in
females.
Distribution. Reunion.
Material examined (1q\ 1 9 syntypes)
Reunion.
Chrysoperla pudica (Navas)
(Figs 149-159)
/Hetnerobius brevicollis Rambur, 1842: 427.
Holotype sex unknown, Mauritius (Marchal)
[not traced]. [Nomen dubium]
Chrysopa pudica Navas, 1914a: 82. Holotype O",
SOUTH AFRICA: Transvaal, Piet Retief
(Crawshay) (BMNH) [examined].
Chrysopa incongrua Navas, 1914c: 97. Holotype,
sex unknown, SENEGAL-NIGER: Koulikoro
(NAVAS COLL.) [type not traced]. Provi-
sional synonym.
Chrysopa nigriciana Navas, 1931b: 119. Holotype
$, NIGERIA: Guezzan & Air, Prince Sixte de
Bourbon, 1929 (MNHN) [examined]. Syn. n.
Chrysopa (Chrysoperla) pudica (Navas); Tjeder,
1966: 397.
Chrysoperla pudica (Navas); Ohm & Holzel,
1982: 160.
Chrysoperla nigricana (Navas); Brooks & Bar-
nard, 1990: 271.
162
S.J. BROOKS
Fore wing (Fig. 149) d\ 8.5-12 mm; 9,
11-13 mm. Head markings variable, unmarked
or sometimes marked with red post-ocular spot;
red spot on gena; some specimens with short,
black lateral stripe on clypeus. Maxillary palps
unmarked. Antennae as long as fore wing.
Pronotum with median yellow longitudinal
stripe; often with red spot in each corner or
medio-lateral spot; prothoracic setae pale, long
or short. Claw (Fig. 150) with large basal dila-
tion, about two-thirds length of claw hook
(1.57-1.76: 1; mean 1.7). Fore wing with black
gradates; crossveins in basal half of wing black at
each end; 3-6 inner gradates, 4-9 outer gradates;
im short, not meeting 1st Rs crossvein or apex of
im meets crossvein; costal setae short (0.6 mm),
inclined towards wing apex.
Genitalias (Figs 152-156). Apodeme of terg-
ite 9 short, not extending beyond callus cercus;
callus narrow, ovate; 26-35 trichobothria; lip of
sternite 8+9 broad rounded; sternite 8+9 with
subapical ventral swelling. Spinellae numerous,
6-8 spines arranged on tubercle. Gonosetae
numerous, evenly distributed on gonosaccus.
Arcessus lacking dorsal striations; apical mem-
brane short, rounded, abruptly curved, some-
times reflexed. Entoprocessus short, narrow.
Lateral plates of gonarcus broadly reniform;
gonarcus arch broad. Acumen of tignum long,
narrow, swollen apically.
Genitalia? (Figs 157-159). Callus cercus
ovate; 37 trichobothria. Subgenitale heart-
shaped with broad basal extension. Spermatheca
with deep ventral impression; vela short; duct
long.
Remarks. Holzel & Ohm (1991; 1992) identi-
fied a small Chrysoperla species from Mauritius
as Chrysoperla brevicollis (Rambur). I have
examined this material and it is the same as C.
pudica. Unfortunately, the type of brevicollis is
missing so the identity of this species must be
gleaned from the original description. Rambur's
(1842) description is brief and in my opinion it is
impossible to say with certainty the genus, let
alone the species, referred to in this description.
One of the most characteristic features of the
Mauritian Chrysoperla species is its black gradate
crossveins. However, in Rambur's description
the venation is said to be very pale yellowish
green and no mention is made of dark veins. In
view of this I think that it is preferrable to treat
Hemerobius brevicollis as nomen dubium rather
than as a senior synonym of Chrysoperla pudica.
Holzel & Ohm (1992) treat the populations of
pudica in Mauritius and mainland Africa as dis-
tinct species. However, I was unable to find any
consistent character to separate the two taxa. In
general specimens of pudica from Mauritius (fore
wing length 9.5-10.5 mm) are smaller than those
from the mainland. However, the smallest speci-
men that I examined during this study was a male
from Nigeria with a fore wing length of 8.5 mm.
Specimens of pudica can be distinguished by
the pale maxillary palps, large basal dilation of
the claw and dark gradates. Males possess tuber-
culate spinellae and lack dorsal striations on the
arcessus. C. pudica has not been recorded from
Madagascar (Holzel & Ohm, 1992) but this may
simply be due to under collecting rather than
disjunct distribution. C. mutata McL. is close to
pudica and the species have many characters in
common. However, the basal dilation of pudica
is considerably larger than mutata. In mutata the
basal tooth is less than half the length of the claw
hook but in pudica the basal tooth is two-thirds
the length of the claw hook. Also the angle
between the basal tooth and the claw hook is
greater in mutata than pudica.
I was unable to trace the type of Chrysopa
incongrua Navas but it seems likely that the
species is synonymous with pudica. Navas
(1914c) describes the species as close to congrua
with a black stripe on the lateral clypeus and
yellow palps, typical markings for West African
specimens of pudica. Nevertheless, he does not
mention any darkening of the gradate crossveins
which casts doubt on this interpretation.
Geographic variation. West African speci-
mens are marked with a short black fleck later-
ally on the clypeus. Southern African specimens
are marked with a red post-ocular spot between
the eye and vertex.
Distribution. Throughout Afrotropics.
Material examined (17 d\ 16 $) Ascension
I., Mozambique, Namibia, Nigeria, South
Africa, St Helena, Senegal, Zambia, Zimbabwe.
Chrysoperla mutata (McLachlan)
(Figs 160-170)
Chrysopa mutata McLachlan, 1898: 167. Lecto-
type cf, ALGERIA: Biskra, 30.iii.1895
(Eaton) [Lectotype designated by Kimmins,
1940: 447] (BMNH) [examined].
Chrysopa nepia Navas, 1911: 266. Holotype Q,
ALGERIA (MNHN) [examined]. Syn. n.
Chrysopa libera Navas, 1928: 463. LECTOTYPE
Cf [here designated], LIBYA: Cyrenaica,
Oasis Giarabub, 16.iii.1927 (Canfalonieri)
(MCSN) [examined]. Syn. n.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
163
Chrysopa quettana Navas, 1931a: 83. Lectotype
^.PAKISTAN: Baluchistan, Quetta,
21. v. 1930 (MNHN) [Lectotype designated by
Leraut, 1991: 79.] [examined]. [Synonymized
with carnea by Aspock et «/., 1980: 268.] Syn.
n.
Chrysopa phaeocephala Navas, 1931b: 120.
Holotype 0\ TUNISIA: Bou Medma
(Dumont) (MNHN) [examined]. Syn. n.
Chrysopa grazianii Navas, 1932: 418. LECTO-
TYPE Cf [here designated], LIBYA: Cufra,
v.1931 (MCSN). [examined]. Syn. n.
Chrysopa expurgata Tjeder, 1949: 83. Holotype
Cf, ISRAEL: Kinneret, -200m, lO.v.1936 (Pal-
moni) (PALMONI COLL., ISRAEL) [not
examined]. [Synonymized by Aspock et at.,
1980: 270.]
Chrysopa (Chrysoperla) expurgata Tjeder;
Tjeder, 1966: 391.
Chrysopa (Chrysoperla) tnutata McLachlan; H61-
zel, 1967: 35.
Anisochrysa (Chrysoperla) mutata (McLachlan);
Holzel, 1970:51.
Anisochrysa (Chrysoperla) concinna Holzel,
1974: 44. Holotype d\ Iran: Bandarabbas,
3-10.iv.1972 (Ressl) (ASPOCK COLL.)
[examined]. Syn. n.
Chrysoperla mutata (McLachlan); Canard &
Laudeho, 1978: 71.
Chrysoperla concinna (Holzel); Holzel, 1980:
169.
Chrysoperla nepia (Navas); Brooks & Barnard,
1990:271.
Chrysoperla phaeocephala (Navas); Brooks &
Barnard, 1990: 271.
Fore wing (Fig. 160) cf, 9-10.5 mm; $,
9.5-12.5 mm. Head marked or unmarked on
gena with faint red spot; short narrow black
stripe on lateral clypeus. Maxillary palp
unmarked. Antennae as long as fore wing.
Pronotum marked with longitudinal median yel-
low band; sometimes with dorso-lateral median
red/brown spot or greyish antero-lateral spot;
prothoracic setae long or short, pale. Basal dila-
tion of claw about half length of claw hook
(1.71-2.64: 1; mean 2.29) (Fig. 161-163). Fore
wing crossveins dark at each end, except mar-
ginal veins at wing margin unmarked; gradates
black; tympanal organ often marked with black
spot; costal setae short (0.6 mm), inclined
towards wing apex; 1-5 inner gradates; 2-6 outer
gradates; cell im variable in length.
Genitalias (Figs 164-167). Lip of sternite 8+9
small, rounded; sternite 8+9 with small subapical
swelling. Apodeme of tergite 9 short, not extend-
ing beyond callus cercus; callus cercus narrow,
ovate; 23-33 trichobothria. Spinellae numerous,
each tubercle bearing 5-7 spines. Gonosetae
numerous, about 25 in lateral group. Arcessus
lacking dorsal striations; apical membrane long,
bent ventrally at 45-90°, tip pointed. Entoproces-
sus narrow, crescentic. Gonarcus with lateral
plate broad, reniform; arch broad. Acumen of
tignum long, narrow or swollen apically.
Genitalias (Figs 168-170). Callus cercus
rounded; 33 trichobothria. Subgenitale heart-
shaped. Spermatheca with moderate ventral
impression; vela short; duct long.
Remarks C. mutata is most similar to puclica
but can be distinguished by the shape of the claw.
In mutata the basal dilation of the claw is about
half the claw hook length, the posterior margin is
relatively straight and the angle between the
basal tooth and the claw hook is relatively large
whereas in puclica the basal dilation is more than
half the claw hook length, the posterior margin is
curved and the angle between the basal tooth and
the claw hook is less than in mutata.
Distribution From north Africa through Ara-
bia and the Middle East to Pakistan and northern
India.
Material examined (26cf, 24$) Algeria,
Egypt, India (Punjab, Rajastan), Iran, Iraq,
Kuwait, Mauritania, Saudi Arabia, Spain, Tuni-
sia.
Chrysoperla nigrinervis sp. n.
(Figs 171-178)
Fore wing (Fig. 171) d\ 9 mm; 9< 10.5 mm.
Head marked with red spot on gena; long, broad
black stripe on lateral clypeus. Maxillary palps
unmarked. Antennae slightly longer than fore
wing. Pronotum marked with longitudinal
median yellow band; greyish lateral stripe; pro-
thoracic setae short, pale. Claw (Fig. 178) with
very small basal dilation, less than one-quarter
length of claw hook (5.8: 1). Fore wing costal
setae short (0.6 mm), inclined towards wing
apex; crossveins entirely black; longitudinal
veins black adjacent to crossveins; tympanal
organ marked with black spot; gradates black;
3-4 inner gradates, 5-6 outer gradates; cell im
short not reaching 1st Rs crossvein or meets at
apex. Abdominal tergites marked with large
black spot on posterior margin.
Genitalias (Figs 172-174). Lip of sternite 8+9
small, rounded; sternite 8+9 with small subapical
swelling. Apodeme of tergite 9 short, not extend-
164
S.J. BROOKS
ing beyond callus cercus; callus cercus narrow,
ovate; 31-32 trichobothria. Spinellae numerous,
each tubercle bearing 5-7 spines. Gonosetae
numerous, about 20 in lateral group. Arcessus
lacking dorsal striations; apical membrane long,
bent ventrally at 45°, tip pointed. Entoprocess
small, narrow, crescentic. Gonarcus with broad
lateral reniform plate; arch broad. Acumen of
tignum long, narrow, slightly swollen at apex.
Genitalia9 (Figs 175-177). Callus cercus
ovate; 37 trichobothria. Subgenitale with long
narrow basal extension. Spermatheca with mod-
erate ventral impression; duct long; vela long.
Remarks. The male genitalia of nigrinervis are
apparently indistinguishable from mutata but
there are sufficient external characters to regard
specimens from the Canary Islands as a distinct
species. C. nigrinervis can be distinguished from
mutata and the closely related pudica by the
more extensively blackened venation, head, pro-
thoracic and abdominal markings together with
the small basal dilation of the claw. In addition,
in females of nigrinervis the subgenitale has a
narrower basal lobe than specimens of mutata
and pudica which have a heart-shaped subgeni-
tale.
Distribution. Canary Islands.
Material examined (2d\ 3$) Holotype d\
Canary Islands: Gran Canada, Maspalomas,
14.viii.1966 (Guichard & Ward) (BMNH)
Paratypes lo", same data as holotype; 3$, same
data as holotype, 17-28. vi. 1966 (BMNH)
Chrysoperla exul (McLachlan)
(Figs 179-187)
Chrysopa exul McLachlan, 1869: 23. Holotype
0\ ST HELENA (Melliss) (BMNH) [exam-
ined].
Chrysopa wollastoni Navas, 1914a: 80. Holotype
(apex of abdomen missing), ST HELENA,
1877 (Wollaston) (BMNH) [examined]. [Syn-
onymized by Kimmins, 1940: 446].
Chrysoperla exul (McLachlan); Holzel, 1989b:
176.
Fore wing (Fig. 179) d\ 12-13 mm, $,
13-14.5 mm. Head marked with red spot on
outer angle of frons; red lateral spot on clypeus;
red spot on lateral vertex; red spot between base
of antennae; scape with faint lateral red stripe;
maxillary palps black dorsally. Antennae longer
than fore wing. Pronotum marked with brown
spot in each corner and red cento-median spot;
median yellow longitudinal band; prothoracic
setae short, pale. Claw (Fig. 180) with large basal
dilation, about two-thirds length of claw hook
(1.74: 1). Fore wing quite broad (length: breadth
= 2.7-3.0: 1); costal setae short (0.4-0.6 mm);
venation entirely green; 6-8 inner gradates, 7-9
outer gradates; im variable in length.
Genitalias (Figs 181-184). Lip of sternite 8+9
short, broad; callus cercus broad ovate; 33 tri-
chobothria; apodeme of tergite 9 branched,
extending only a short way beyond callus cercus.
Spinellae present, 6-7 spines on numerous small
tubercles. Gonosetae numerous. Entoprocessus
small, crescentic. Arcessus straight, extreme
apex curved at 45°; lacking dorsal striations;
apical membrane of arcessus long, narrow, trun-
cate apically. Gonarcus with broad reniform lat-
eral plate; arch narrow. Acumen of tignum long,
narrow.
Genitalia9 (Figs 185-187). Callus cercus
ovate; 38 trichobothria. Subgenitale with narrow
basal extension. Spermatheca with shallow ven-
tral impression; vela short; duct long.
Remarks. C. exul may be distinguished from
other species of Chrysoperla by the characteristic
head and pronotal markings, in particular the
brown spot in each corner of the pronotum and
the red spots on the vertex and between the
antennae.
The possession of synapomorphies in exul and
congrua suggests a close relationship between
these two species. In both these species the
gonarcus arch is narrow and the spinellae are
tuberculous.
Distribution. St Helena.
Material examined (6d\ 4$) St Helena.
Chrysoperla congrua (Walker)
(Figs 188-195)
Chrysopa congrua Walker, 1853: 238. Holotype
(abdomen missing), WEST AFRICA
(BMNH) [examined].
Chrysopa concolor Walker, 1853: 239. Holotype
(abdomen missing), ZAIRE (BMNH) [exam-
ined]. [Synonymized by Tjeder, 1966: 392.]
Chrysopa otalatis Banks, 1910a: 102. Holotype
$, AUSTRALIA: Queensland, Brisbane,
vi.1904 (MCZ) [not available for examina-
tion]. Syn. n.
Chrysopa bequaerti Navas, 1912: 409. Lectotype
d\ ZAIRE: Kongolo, 9.ii.l911 (Bequaert)
[lectotype designation by Tjeder, 1966: 393.]
(MRAC) [examined]. [Synonymized by
Esben-Petersen, 1928a: 218.]
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
165
Chrvsopa lemoulti Lacroix, 1923: 119. Holotype
Cf, NEW CALEDONIA (MNHN) [exam-
ined]. [Synonymized with otalatis by Esben-
Petersen, 1928b: 103.] Syn. n.
Chrvsopa clypealis Navas, 1929b: 103. Holotype
$, ZAIRE: Haut-Uele, Yebo Moto, Abmiva.
1925 {Burgeon) (MRAC) [examined]. Syn. n.
Chrysopa solaria Navas, 1930: 411. Holotype 0\
EAST AFRICA: Marangu (ZMH)
[destroyed]. [Synonymized by Holzel, 1989b:
166.]
Chrysopa bequaerti var. occipitalis Navas, 1931c:
127. Holotype $, ZAIRE (MRAC) [exam-
ined]. Syn. n.
Chrysopa congrua var. rubella Navas, 1931c:
127. Holotype $, ZAIRE (MRAC) [exam-
ined]. Syn. n.
Chrvsopa acutella Navas, 1933a: 209. Holotype
Cf .MADAGASCAR: Majunga (MNHN)
[examined]. [Synonymized by Holzel, 1989b:
166.]
Chrysopa (Chrysoperla) congrua (Walker);
Tjeder, 1966:392.
Chrysoperla congrua (Walker); Semeria & Qui-
lici, 1986: 110.
Chrysoperla otalatis (Banks); Brooks & Barnard,
1990:271.
Chrysoperla acutella (Navas); Brooks & Bar-
nard, 1990: 271.
Chrvsoperla clypealis (Navas); Brooks & Bar-
nard, 1990:271.
Fore wing (Fig. 188) O" 10-12 mm; 9
12-13.5 mm. Head usually marked with red lat-
eral stripe on gena, clypeus, sometimes extend-
ing to frons; red post-ocular spot often present.
Maxillary palp unmarked or marked black dor-
sally. Antenna about as long as fore wing. Prono-
tum with longitudinal median yellow band and
often with red or greyish spot in anterior corner;
prothoracic setae long, pale (short in Easter I.
specimens). Claw (Fig. 189) with small basal
dilation, less than one quarter length of claw
hook (4.1-6.25: 1; mean 5.05). Fore wing narrow
(length: breadth = 3.0-4.2: 1), pointed apically;
5-8 inner gradates, 5-9 outer gradates; im very
short; costal setae moderately long
(0.8-1.0 mm), inclined; veins green except cl
and c2 crossveins maybe darkened; 1st Rs cross-
vein at right-angles to Rs.
Genitalias (Figs 190-192). Lip of sternite 8+9
short, narrow, downcurved. Callus cercus broad,
ovate; 24-33 trichobothria; apodeme of tergite 9
extends beyond callus cercus. Spinellae sparse,
4-6 spines arranged on small tubercles. Gonose-
tae numerous, evenly distributed across gonosac-
cus. Arcessus narrow, weakly curved, lacking
dorsal striations, apex blunt and slightly swollen.
Entoprocessus very small, short and narrow.
Gonarcus with narrow arch and narrow, wedge-
shaped lateral plates. Acumen of tignum short,
bulbous.
Genitalia9 (Figs 193-195). Callus cercus
rounded. Basal extension of subgenitale usually
long and narrow. Spermatheca small with, deep
ventral impression; vela short; duct long.
Remarks. C. congrua is characterized by pos-
sessing green gradates, a small basal dilation of
the claw and narrow fore wings with a short
intramedian cell. A close relationship between
congrua and the Neotropical species of Chrysop-
erla is suggested by the synapomorphous posses-
sion of a swollen apex to the arcessus and
bulbous acumen (cf. C. externa).
From New's (1980) description of Chrysopa
itatotis Banks, evidently based on the female
holotype. it is apparent that the species is refer-
able to Plesiochrysa (comb. n.). However, the
figure he gives of the male genitalia are those of a
Chrysoperla species. On examination of the eight
specimens New (1980) identified as C. italotis,
from the ANIC, it is apparent that in fact these
are misidentfied specimens of Chrysoperla con-
grua.
Geographical variation. Most material
examined from Australia, New Caledonia and
New Hebrides had the maxillary palps
unmarked. In specimens from Samoa, Easter
Island and West Malaysia the palps were always
black dorsally. The palps were variable in speci-
mens from other regions. All the Sri Lankan
specimens were collected between 4,000 and
7,000 feet.
Distribution. Throughout Australasia from
the Pacific Islands to Sri Lanka (absent from
India), throughout the Afrotropics and recorded
from Oman on the Arabian Peninsular (Holzel &
Ohm, 1992)
Material examined (117o\ 196$). Afrotro-
pics: Angola, Botswana, Ethiopia, Fernando Po,
Gabon, Ghana, Ivory Coast, Kenya, Madagas-
car, Malawi, Mozambique, Nigeria, Prinicipi
Island, Ruanda, South Africa, Tanzania,
Uganda, Urundi, Zaire, Zambia, Zimbabwe.
Oriental: Sri Lanka, Thailand, West Malaysia.
Australasian: Australia, New Caledonia, New
Hebrides, Papua New Guinea. Pacific: Easter I.,
Fiji, Rapa I., Samoa.
166
S.J. BROOKS
Chrysoperla dozieri (Smith)
(Figs 196-202)
Chrysopa dozieri Smith, 1931: 808. Holotype G\
HAITI: Port au Prince, 29. ii. 1929 (Smith)
(MCZ) [examined].
Chrysoperla dozieri (Smith); Brooks & Barnard,
1990: 271.
Fore wing (Fig. 196) 0\ 10.5-12 mm; §, not
seen. Head marked with red L-shaped stripe
across frontal suture; red stripe on gena and
lateral clypeus; narrow medio-lateral red spot on
front of vertex. Maxillary palps marked black
dorsally. Antennae as long as fore wing. Prono-
tum marked with longitudinal median yellow
band; medio-lateral black spot and usually with
red stripe in anterior corner; prothoracic setae
short, pale. Claw (Fig. 197) with large basal
dilation, about half length of claw hook (2.08: 1).
Fore wing pointed at apex; costal setae short
(0.4—0.6 mm), inclined towards wing apex; 5-6
inner gradates, 6-8 outer gradates; gradates
black; other crossveins black each end; cell im
long, 1st Rs crossvein meets im at apex or
sub-apically; 1st Rs crossvein straight or oblique.
GenitaliacT (Figs 198-202). Lip of sternite 8+9
broad, rounded. Apodeme of tergite 9 forked
above and below callus, short; callus cercus
broad, ovate; 29 trichobothria. Spinellae few,
tuberculate. Gonosetae numerous, in lateral
group. Arcessus broad, straight; dorsal striations
absent; membranous tip spade-like (narrows sub-
apically, then abruptly widens at apex), truncate
at apex, curved ventrad at 90°. Entoprocessus
not visible. Gonarcus lateral plates broadly trian-
gular; arch broad. Acumen of tignum narrow,
hardly swollen apically.
Remarks. This species can be readily distin-
guished from other species in the genus by the
presence of a pair of narrow red spots on the
vertex. Some Cuban specimens of exotera may
also have a pair of red spots on the front of the
vertex but they are broader than those of dozieri.
Unlike dozieri, specimens of exotera do not have
a red stripe below the fronto-clypeal suture. The
spade-like apex of the arcessus in the male geni-
talia is also distinctive. C. dozieri differs from
other Chrysoperla species in the rufilabris-group
in plesiomorphically retaining a broad gonarcus
arch and a broad arcessus. The species also has
fewer spinellae tubercles than other rufilabris-
group species. C. dozieri is endemic to Haiti and
this geographical isolation may account for its
apparent phylogenetic isolation.
Distribution. Haiti.
Material examined (2d\ 2 examples with
abdomens missing) Haiti.
Chrysoperla defreitasi sp. n.
(Figs 203-208)
Fore wing cf (Fig. 203), 11 mm; $ unknown.
Head marked with broad red stripe on gena,
lateral clypeus and lateral labrum; narrow red
stripe on lateral fronto-clypeal suture. Maxillary
palps marked black dorsally. Pronotum with yel-
low longitudinal median stripe and narrow
brownish red lateral stripe; prothoracic setae
long, pale. Claw (Fig. 208) with large basal
dilation, about two-thirds length of claw hook
(1.68: 1). Fore wing relatively narrow (length:
breadth = 3: 1); venation mostly green with
black gradates, basal costal crossveins entirely
black, black spot at each end of radial and Rs
crossveins, c,/c 2 crossvein entirely black; mar-
ginal branches of veins Cu 2 , 1A and 2 A entirely
black; apex of fore wing pointed; costal setae
short (0.6 mm), inclined towards wing apex; im
short, not reaching 1st Rs crossvein; 5 inner
gradates, 7 outer gradates.
GenitaliacT (Figs 204-207). Dorsal apodeme of
tergite 9 long; callus cercus narrow ovate; 28-29
trichobothria; lip of sternite 8 + 9 short, broad.
Spinellae present, 29 large tubercles with 7-8
spicules. Gonosetae numerous 50+ in lateral
group. Arcessus broad, tapering subapically
before widening to form spatulate apex; dorsal
striations absent. Entoprocessus narrow, cres-
centic. Gonarcus with broad, wedge-shaped lat-
eral plates; arch relatively broad, truncate
medially with bilobed median projection. Acu-
men of tignum long, narrow, slightly swollen at
apex.
Remarks. C. defreitasi closely resembles C.
externa and C. exotera. From C. exotera, C.
defreitasi cannot be reliably distinguished on
external characters although the claw hook is
larger in defreitasi than in exotera. However, the
male genitalia are quite different. In C. defreitasi
the arcessus is broad with a spatulate apex
whereas in C. exotera the arcessus is narrow and
truncate at the apex.
C. defreitasi may be distinguished from externa
by the black gradate crossveins, which are green
in externa, and the basal dilation of the claw,
which is considerably larger in defreitasi than
externa. Certain apomorphic charcters in the
male genitalia suggest that C. defreitasi may be
closely related to C. dozieri, from Haiti. In both
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
167
species the unstriated arcessus is broader than is
typical in neotropical species of the genus, the
apex of the arcessus is spatulate and the gonarcus
arch is relatively broad. Although the entopro-
cessus are narrow in C. defreitasi, they are larger
than other Chrysoperla species in species-group
4.2 (see p. 143).
Distribution. Brazil.
Material examined (Id") Holotype cf, Bra-
zil: Jaquaritingas, 15. xi. 1992 (de Freitas) (USP).
Chrysoperla asoralis (Banks)
(Figs 209-217)
Chrysopa asoralis Banks, 1915: 627. Holotype
0\ PERU: Chosica, 2800', 7.vi. (Parish)
(MCZ) [examined].
Chrysopa oscillans Navas, 1922a: 260. LECTO-
TYPE Cohere designated], PERU: Arequipa,
1912 (Escomel) (MNHN) [examined]. Syn. n.
Chrysopa lindana Navas, 1924b: 111. Holotype
$, PERU: on the Chilean frontier, 1921 (Mar-
tin) (MNHN) [examined]. Syn. n.
Chrysopa arequipae Navas, 1929a: 18. Holotype
$,PERU: Arequipa, 1922 (Escomel)
(MNHN) [examined]. Syn. n.
Chrysoperla asoralis (Banks); Brooks & Bar-
nard, 1990:271.
Fore wing (Fig. 209) d\ 10-14 mm, $.
12-16 mm. Head marked with red stripe on
gena, lateral clypeus and frons, occasionally with
vertex red dorsally; red post-ocular spot. Maxil-
lary palps black dorsally. Antennae as long as
fore wing. Pronotum with median longitudinal
yellow stripe; red spot in anterior corner; protho-
racic setae short, pale. Claw (Fig. 210) with large
basal dilation, over half length of claw hook
(1.83-1.87: 1; mean 1.85). Fore wing pointed at
apex; costal setae short (0.6-0.8 mm), inclined
towards wing apex; cell im variable in length; 1st
Rs crossvein meets Rs at right-angles; 5-8 inner
gradates, 6-9 outer gradates; gradates black;
other crossveins dark at each end, except Psm
crossveins unmarked.
Genitalias (Figs 214-217). Lip of sternite 8 + 9
broad, rounded. Dorsal apodeme of tergite 9
long, extending well-beyond callus cercus; callus
cercus narrow, ovate; 20-41 trichobothria.
Spinellae numerous, each tubercle with 4-5 long
spines. Gonosetae numerous, 50+ in lateral
group. Arcessus narrow, arcuate, occasionally
recurved apically; dorsal striations absent; apical
membrane long, pointed at tip. Entoprocessus
minute, slender, crescentic. Gonarcus arch nar-
row; narrow wedge-like lateral plate. Acumen of
tignum long, swollen at apex.
Genitalia9 (Figs 211-213). Callus cercus
rounded; 32 trichobothria. Subgenitale with nar-
row basal extension. Spermatheca small with
moderate ventral impression; vela moderate;
duct long.
Remarks. C asoralis most closely resembles
exotera and ruflabris, both of which have black
gradate crossveins and black maxillary palps.
From exotera, asoralis can be distinguished by
the red post-ocular spot, which is absent in
exotera, and the apical membrane of the arcessus
which is pointed at the tip in males of asoralis but
truncate and slightly swollen in males of exotera.
C. rufilabris also differs from asoralis in the
shape of the apex of the arcessus which is much
longer and is abruptly narrowed in the former
species. In addition, the acumen is shorter and
broader in rufilabris than asoralis. The spinellae
are unusually long in asoralis.
The type of lindana differs from other speci-
mens of asoralis in having a longer fore wing
(16 mm, which was 2 mm longer than the next
largest specimen) and the vertex marked red with
a narrow median interruption. However, I con-
sider these differences to be due to individual
variation and not of taxonomic significance.
Distribution. Neotropics.
Material examined (20cf, 149) Argentina,
Bermuda. Guatemala, Peru, Venezuela.
Chrysoperla exotera (Navas)
(Figs 218-226)
Chrysopa exotera Navas, 1914a: 87. Holotype $,
GUATEMALA: Duenas, Capetillo (Cham-
pion) (BMNH) [examined].
Chrysopa extranea Navas, 1922b: 170. Holotype
Cf .CUBA (MCZ) [examined]. Syn n.
Chrysopa estradai Navas, 1924a: 334. Holotype,
CUBA: Habana, Rio Almendares,
23-26.X.1923 (Cervera) (HAVANA) [not
examined]. Syn. n.
Chrysopa torrei Navas, 1924a: 335. Holotype cf ,
CUBA: Habana, Rio Almendares, 8.ix.l923
(Cervera) (MCZ) [examined]. Syn. n.
Chrysopa barbouri Navas, 1924a: 335. Holotype
sex unknown (abdomen missing), CUBA:
Habana, Rio Almendares, 8.xii.l923 (Cervera)
(MCZ) [examined]. Syn. n.
Chrysopa tenera Navas, 1924a: 336. Holotype $,
CUBA: Habana, Rio Almendares, 8.xii.l923
(MCZ) [examined]. Syn. n.
168
S.J. BROOKS
Chrysopa exterior Navas, 1925: 190. [Replace-
ment namefor C. extranea Navas].
Chrysopa cubensis Navas, 1927a: 75. [Unneces-
sary replacement name for C. extranea Navas].
Chrysopa incisa Banks, 1948: 163. Holotype £,
MEXICO: Chihuahua, Ciudad Juarez,
4.vi.l927 (Dampf) (MCZ) [not examined, not
traced in MCZ]. Syn. n.
Chrysopa quadornia Banks, 1948: 164. Holotype
$, MEXICO: Acapulco, 19.xii.1924 {Dampf)
(MCZ) [not examined, not traced in MCZ].
Syn. n.
Chrysopa castalia Banks, 1948: 165. Holotype $,
MEXICO: Tamaulipas, Santa Engracia,
3.ii.l936 {Dampf) (MCZ) [not examined, not
traced in MCZ]. Syn. n.
Chrysoperla exotera (Navas); Brooks & Barnard,
1990: 271.
Fore wing (Fig. 218) cf, 13 mm; 9,
11.5-13.5 mm. Head marked with red spot on
gena, lateral clypeus and lateral frons; anterior
margin of vertex sometimes with red stripe or
paired spots; entire head occasionally suffused
red. Maxillary palps black dorsally. Antennae
shorter than fore wing. Yellow longitudinal band
on pronotum present or absent; occasionally with
red spot in anterior corner; prothoracic setae
long, pale. Claw (Fig. 219) with large basal
dilation, about half claw hook length (2.07: 1).
Fore wing apex pointed; costal setae short
(0.6-0.8 mm), inclined towards wing apex; gra-
dates black or greyish; crossveins black at each
end except at wing margins; 5-7 inner gradates,
7-9 outer gradates; cell im short, apex not reach-
ing 1st Rs crossvein; 1st Rs crossvein straight or
oblique.
Genitalias (Figs 220-223). Lip of sternite 8+9
broad, rounded. Dorsal apodeme of tergite 9
short, forked above or below callus cercus; callus
cercus broad, ovate; 40 trichobothria. Spinellae
numerous, about 8 spines on each tubercle.
Gonosetae numerous, arranged in lateral group.
Arcessus narrow, arcuate; dorsal striations
absent; apical membrane short, tip broadened,
truncate. Entoprocessus minute, narrow, cres-
centic. Gonarcus with lateral plates narrow, reni-
form or wedge-shaped; arch narrow. Acumen of
tignum short, broad, swollen at apex.
Genitalia 9 (Figs 224-226). Callus cercus
rounded; 32 trichobothria. Subgenitale with nar-
row basal extension. Spermatheca narrow with
deep ventral impression; vela long; duct moder-
ate.
Remarks. This species is very similar to
comanche but can be distinguished by the black
gradate crossveins which are green in comanche.
In addition, the apex of the arcessus and acumen
in exotera is swollen, whereas in comanche they
are narrow.
Distribution. Central America and Cuba.
Material examined (lcf, 4$) Cuba, Costa
Rica, Guatemala, Mexico.
Chrysoperla externa (Hagen)
(Figs 227-236)
Chrysopa externa Hagen, 1861: 221. Syntypes,
USA: Washington, California; Mexico (MCZ)
[not traced].
Chrysopa lanata Banks, 1910b: 154. Holotype
CT, ARGENTINA: Mendoza {Jensen-Haarup)
(MCZ) [examined]. [Synonymized by Adams,
1962: 222.]
Chrysopa graciana Navas, 1919: 301. Holotype,
ARGENTINA: Alta Gracia, ii.1918 {Bruch)
(LP) [not examined]. [Synonymized with
argentina Navas by Adams, 1975: 169. Synony-
mized with externa by Adams & Penny, 1987:
421.]
Chrysoperla externa (Hagen); Adams, 1983: 43.
Chrysoperla externa cocosensis Adams, 1983: 42.
Holotype Q\ COSTA RICA: Cocos I., Wafer
Bay, 9. hi. 1980 {Sherry & Werner) (LACM)
[examined]. Syn. n.
Fore wing (Fig. 227) Q\ 10.5-11.5 mm; 9,
12.5 mm. Head marked with broad red stripe on
gena and lateral clypeus; red post-ocular stripe.
Maxillary palps black dorsally. Antennae about
same length as fore wing. Pronotum with yellow
longitudinal median band; usually with reddish
spot in anterior corner; prothoracic setae long or
short, pale or black. Claw (Fig. 228-229) with
small basal dilation, less than half claw hook
length (2.4-2.54: 1; mean 2.48). Fore wing nar-
row (length: breadth = 3.1-3.5: 1), apex
pointed; costal setae short (0.6-1.0 mm),
inclined towards wing apex; 5-7 inner gradates,
6-8 outer gradates; gradates green; crossveins in
basal half of wing often darkened at each end;
cell im very short, not reaching 1st Rs crossvein;
1st Rs crossvein meets Rs at right angles.
Genitalias (Figs 230-233). Lip of sternite 8+9
rounded. Dorsal apodeme of tergite 9 long,
forked below callus; callus cercus narrow, ovate;
28-32 trichobothria. Spinellae numerous, each
tubercle with 5-6 spines. Gonosetae numerous,
positioned centrally on gonosaccus. Arcessus
without dorsal striations; membranous apex of
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
169
arcessus short, bulbous, curved ventrally at 45°.
Entoprocessus small triangular, often not visible.
Gonarcus with lateral plates narrow, wedge-
shaped; arch narrow. Acumen of tignum long,
slightly swollen apically.
Genitalia9 (Figs 234-236). Callus cercus
ovate; 32 trichobothria. Subgenitale with broad,
rounded basal extension. Spermatheca narrow;
ventral impression deep; vela short; duct long.
Remarks. C. externa is similar in appearance to
comanche but the basal dilation of the claw is
smaller in externa and the apex of the arcessus is
narrow in comanche but swollen in externa. C.
externa appears to be closely related to congrua.
since the male genitalia are very similar and the
apex of the arcessus is swollen in both species.
However, although the apex of the acumen is
swollen in both externa and congrua, the neck is
longer in externa. In addition the basal dilation of
the claw is noticeably larger in externa than
congrua.
I have examined a female specimen deposited
in the MCZ which has been identified as C.
externa Hagen by Banks (USA: Florida, Winter
Park, 31.x., Davis). This specimen bears a purple
label, type written 'neotype'. I have not been
able to trace any reference to this specimen being
nominated a neotype so this designation has no
validity. The specimen has been correctly identi-
fied but since there is no confusion regarding the
identity of C. externa and considering that the
specimen was not collected in the same region as
the original type series I see no merit in formally
recognising this specimen as a neotype.
Adams (1983) distinguished C. e. cocosensis
from C. e. externa on the basis of the arcessus
being sclerotised medially in the former taxon
but sclerotised laterally with a median membra-
nous area in the latter. However, examination of
the holotype of C. e. cocosensis revealed that
Adams had misinterpretted the structure of the
arcessus and that, in fact, the lateral sclerotised
strips are separated by a membranous median
area. The arcessus of other males examined from
Cocos Island were also indistinguishable from
typical examples of C. externa.
Geographical variation. In Mexican speci-
mens the post-ocular spot is absent, the lateral
frons marked with red stripe, the basal dilation of
the claw larger and the intramedian cell longer
than typical specimens.
Distribution. Throughout Neotropics, extend-
ing northwards to Florida and South Carolina
(Adams, 1962).
Material examined (27d\ 48$) Argentina,
Belize, Brazil, Costa Rica (Cocos Island), Guate-
mala, Mexico, Peru, Uruguay, USA (Florida),
Venezuela, West Indies (Bahamas, Barbados,
Cayman, Trinidad).
Chrysoperla galapagoensis (Banks)
(Figs 237-245)
Chrysopa galapagoensis Banks, 1924: 179. Holo-
type, ECUADOR: Galapagos Is, South Sey-
mour, 23. iv. 1923 (Beebe) (MCZ) [not
examined].
Chrysoperla galapagoensis (Banks); Brooks &
Barnard, 1990:271.
Fore wing (Fig. 237) cf, 9-10.5 mm; $,
10.5 mm. Head marked with black stripe on gena
and lateral clypeus; red stripe on lateral frons,
lateral labrum and dorso-lateral clypeus; red
post-ocular stripe present or absent. Maxillary
palps black dorsally. Antennae shorter than fore
wing. Pronotum usually marked with narrow
black median stripe and latero-median black
spot, yellow longitudinal stripe absent; protho-
racic setae long, black or pale. Claw (Fig. 238)
with small basal dilation, less than half length of
claw hook (2.73: 1). Fore wing pointed apically;
costal setae short (0.6-0.8), inclined towards
wing apex; 4-6 inner gradates, 5-6 outer gra-
dates; gradates black; crossveins entirely black,
except Psc crossveins in apical half of wing; cell
im short, not quite meeting 1st Rs crossvein or
crossvein meets apex of cell; 1st Rs crossvein
oblique or at right-angles to Rs.
Genitalias (Figs 239-241). Lip of sternite 8+9
narrow, rounded. Apodeme of tergite 9 short,
branched below callus cercus; callus cercus nar-
row, ovate; 23-26 trichobothria. Spinellae few,
6-12 small tubercles bearing about 4 spines.
Gonosetae numerous. Arcessus narrow, gently
curved; lacking dorsal striations; apex rounded
and slightly expanded. Entoprocessus not visible.
Gonarcus with lateral plates narrow, wedge-
shaped; arch narrow. Acumen of tignum short,
slightly swollen apically.
GENITALIA9 (Figs 242-245). Callus cercus
broadly ovate; 28 trichobothria. Subgenitale with
short, narrow basal extension. Spermatheca with
shallow ventral impression; vela short; duct
short.
Remarks. C. galapagoensis is distinctive
amongst Chrysoperla species because of the
extensive black venation, prominent black lateral
stripe on the head and narrow black median
170
S.J. BROOKS
stripe on the pronotum. The male genitalia are
also distinctive since the spinellae are few and
small. The arcessus is slightly swollen apically
which is reminiscent of externa but the tip of the
arcessus is more bulbous in externa than galapa-
goensis.
Distribution. Galapagos Islands.
Material examined (5cf, 5$) Galapagos
Islands (Fernandina, Hood, Indefatigable,
Jervis, Santiago, South Seymour, Tower).
Chrysoperla comanche (Banks)
(Figs 246-255)
Chrysopa comanche Banks, 1938: 119. Holotype
$, USA: Texas, Laredo (MCZ) [examined].
Chrysopa sperryae Banks, 1943: 74. Holotype $,
USA: California, Riverside (Sperry) (MCZ)
[not examined]. [Synonymized by Adams,
1956: 45.]
Chrysoperla comanche (Banks); Garland, 1985b:
738.
Fore wing (Fig. 246) cf, 8-11 mm; $,
12-12.5 mm. Head marked with red stripe on
gena and lateral clypeus, usually extending to
lateral frons; red post-ocular spot present or
absent; occasionally head totally unmarked.
Maxillary palps black dorsally. Antennae about
as long as fore wing. Pronotum with yellow
longitudinal median stripe; sometimes with
dorso-median black spot. Claw (Fig. 247) with
large basal dilation, over half length of claw hook
(1.86-2.0: 1; mean 1.93). Fore wing apex
pointed; costal setae short (0.4—0.6 mm),
inclined towards wing apex; Rs crossveins in
stigmatic region sinuate; 3-6 inner gradates, 4—8
outer gradates; gradates green, sometimes inner
gradates greyish; crossveins in basal half of wing
sometimes dark at ends; cell im short, not reach-
ing 1st Rs crossvein; 1st Rs crossvein oblique or
at right-angles to Rs.
Genitalias (Figs 248-252). Lip of sternite 8+9
broad, rounded. Apodeme of tergite 9 long,
forked below callus; callus cercus narrow, ovate;
26-29 trichobothria. Spinellae numerous, 4—5
long spines on tubercle. Gonosetae numerous,
50+ in lateral group. Arcessus narrow, arcuate;
dorsal striations absent; apical membrane short,
truncate or slightly broadening. Entoprocessus
minute, slender crescentic. Gonarcus with nar-
row lateral plates; arch narrow. Tignum with
acumen long, slightly widened apically.
Genitalia9 (Figs 253-255). Callus cercus
rounded; 30 trichobothria. Subgenitale with
long, narrow basal extension. Spermatheca with
moderate ventral impression; duct short; vela
moderate.
Remarks. C. comanche is very similar to
exotera but there are two characters that suggest
they are distinct species. Specimens assignable to
exotera have black gradate crossveins, swollen
acumen and the arcessus is markedly swollen at
the extreme apex. In comanche the gradates are
green, the acumen is narrow and the apex of the
arcessus is narrow or slightly widened. The two
species are also separated geographically, with
exotera occurring in Central America but
comanche being restricted to southwest USA. C.
comanche can be distinguished from most New
World Chrysoperla species by the green gra-
dates. From the carnea-group of species
comanche can be separated by the red stripe on
the gena since the gena is marked brown in the
carnea-group. From externa, comanche differs by
having a large basal dilation of the claw, and
narrow arcessus. In externa the basal dilation of
the claw is distinctly less the half the length of the
claw hook and the arcessus is swollen apically.
Material examined from the Hawaiian Islands
had been previously identified as C. lanata Banks
(Zimmerman, 1957), which is now considered as
a synonym of C. externa (Hagen) (Adams, 1962).
However, all Hawaiian specimens examined in
this study proved to be comanche and there is no
evidence that externa occurs on the Hawaiian
Islands.
Distribution. South-west USA, Hawaii.
Material examined (51d\ 48$) . USA (Ari-
zona, California); Hawaiian Islands (Kauai,
Lanai, Maui, Molokai, Oahu).
Chrysoperla harrisii (Fitch)
(Figs 256-265)
Chrysopa perla Harris, 1841: 197. Holotype,
USA: New England (MCZ) [not examined].
[Homonym of Chrysopa perla L.]
Chrysopa harrisii Fitch, 1855: 794. [New name
for C. perla Harris].
Chrysopa stenostigma Navas, 1914b: 61. Holo-
type (sex unknown), USA: New York, Long
Island, Yaphank, 25.vii.1913 (NAVAS
COLL.) [not traced]. [Synonymized by Bick-
ley & MacLeod, 1956: 193.]
Chrysoperla harrisii (Fitch); Garland, 1985a: 93.
Fore wing (Fig. 256) cf, 12.5 mm; $, 13-14 mm.
Head marked with broad red stripe on gena,
lateral frons and lateral clypeus; broad red post-
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
171
ocular stripe. Maxillary palps black dorsally.
Antennae slightly longer than fore wing. Prono-
tum marked with yellow longitudinal median
band; brownish spot in anterior corner; protho-
racic setae short, black or pale; setal follicles
black. Claw (Fig. 257) with very small basal
dilation, less than one-quarter claw hook length
(8: 1). Fore wing narrow (length: breadth =
3.2-3.4: 1), apex pointed; venation entirely
green; costal setae short (0.8-1.0 mm), inclined
towards wing apex; cell im long, 1st Rs crossvein
meets im at apex or subapically; 1st Rs crossveins
leaves Rs obliquely; 7-9 inner gradates, 9-10
outer gradates.
Genitalias (Figs 258-262). Lip of sternite 8+9
broad, rounded. Dorsal apodeme of tergite 9
long, extending far beyond callus cercus; callus
cercus narrow, ovate; 32 trichobothria. Spinellae
present, numerous tubercles bearing about 7
spines. Gonosetae numerous, arranged in lateral
group. Arcessus without dorsal striations;
strongly curved ventrad; membranous tip bent at
90°, truncate apically. Entoprocessus minute,
crescentic. Lateral plate of gonarcus narrow
wedge-shaped; arch narrow. Acumen of tignum
short, broad, slightly swollen apically.
Genitalia? (Figs 263-265). Callus cercus
broadly ovate; 37 trichobothria. Subgenitale with
long, narrow basal extension. Spermatheca with
shallow ventral impression; vela short; duct
short.
Remarks. C. harrisii is one of the narrow
winged species of Chrysoperla but is character-
ized by the entirely green venation and extremely
small basal dilation of the claw. The male genita-
lia of harrisii are similar to those of comanche
which suggests that the the two species may be
closely related. However, these species may be
differentiated by the head markings, which are
more extensive in harrisii than comanche and the
shape of the acumen, which is broader in harrisii
than comanche.
Distribution. Canada (Garland, 1982); USA:
California, Maryland, Michigan, Virginia (Bram
& Bickley, 1963); Montana, Connecticut;
Mexico.
Material examined (2cf, 3$) . USA: Mon-
tana; Connecticut.
Chrysoperla mexicana sp. n.
(Figs 266-274)
Fore wing (Fig. 266) o", 10.5 mm; $, 11.5 mm.
Head marked with broad red lateral stripe on
gena, frons and clypeus with blackish flecks on
gena and clypeus. Maxillary palp black dorsally.
Antennae about same length as fore wing; outer
margin of scape with apical brown stripe; pedicel
and flagellar segments entirely dark brown.
Pronotum marked with yellow median longitudi-
nal band and brownish red lateral stripe; protho-
racic setae short, black. Claw (Fig. 267)
undilated basally. Fore wing narrow (length:
breadth = 3.4-3.6:1), apex pointed; gradate
crossveins green; costal, radial and Psc cross-
veins black, darker in basal half of wing; veins
7/4,2/4, Cu 2 , base of Rs and Psm at cell M 2 and
im all black; other veins green; costal setae short
(0.6 mm), inclined towards wing apex; cell im
short, 1st Rs meets Psm just beyond apex of im;
1st Rs crossvein leaves Rs at right angles; 5-6
inner gradates, 7-8 outer gradates.
Genitalias (Figs 268-271). Lip of sternite 8+9
broad, rounded. Dorsal apodeme of tergite 9
short, not extending beyond callus cercus. Callus
cercus broad, ovate; 29 trichobothria. Spinellae
present, numerous tubercles with 5-7 spines.
Gonosetae numerous, arranged in lateral group.
Arcessus narrow, without dorsal striations; apex
curved ventrally at 90°; apical membrane moder-
ately long, truncate at tip. Entoprocessus very
narrow, crescentic. Lateral plate of gonarcus
narrow, quadrangular; arch narrow. Acumen of
tignum narrow, swollen at apex.
Genitalia? (Figs 272-274). Callus cercus
broad, ovate; 26 trichobothria. Subgenitale with
long narrow basal lobe. Spermatheca with shal-
low ventral impression (everted in figure); vela
short; duct short.
REMARKS. Chrysoperla mexicana can be readily
distinguished by the black antennae, in all other
species of the genus the antennae are green.
Other distinguishing characters are the broad
brownish lateral stripe on the pronotum, the
undilated base of the claw and the narrow fore
wings with dark venation in the basal half of the
wing.
C. mexicana appears to be closely related to
harrisii since both species share a number of
apomorphies including narrow fore wings, claws
with small basal dilation, truncate apex of arces-
sus, and long basal lobe on the spermatheca.
I am most grateful to Prof. P. A. Adams for
drawing my attention to this species.
Distribution. Mexico.
Material examined (ld\ 1$) Holotype $,
Mexico: Sinaloa, 36.6 miles NE Villa Union,
172
S.J. BROOKS
near La Capilla Detaxte, 2.vii.l982 (SDNHM)
Paratype 9, same data as holotype.
Chrysoperla rufilabris (Burmeister)
(Figs 275-285)
Chrysopa rufilabris Burmeister, 1839: 979. Syn-
types, Mid-USA & MEXICO (ZMH) [type
destroyed].
Chrysopa interrupta Schneider, 1851: 76. Holo-
type d\ USA: Pennsylvania (NMHU) [not
examined]. [Synonymized by Bram & Bickley,
1963: 6.]
Chrysopa attenuata Walker, 1853: 242. Lectotype
d\ USA: Florida, St John's Bluff (Doubleday)
[Lectotype designation by Kimmins, 1940:
447.] (BMNH) [examined]. [Synonymized by
Banks, 1903: 161.]
Chrysopa repleta Walker, 1853: 244. Holotype
(abdomenmissing), USA: Georgia (BMNH)
[examined]. [Synonymized by Banks, 1903:
161.]
Chrysopa novaeboracensis Fitch, 1855: 90. Syn-
types, USA: New York State (MCZ) [not
examined]. [Synonymized by Hagen, 1866:
395.]
Chrysopa tabida Fitch, 1855: 92. USA (MCZ)
[not examined]. [Synonymized by Hagen,
1866: 398].
Chrysopa citri Ashmead, 1880: 13. Holotype,
USA: Florida [not examined]. [Synonymized
by Bickley & MacLeod, 1956: 194.].
Chrysopa medialis Banks, 1903: 154. Holotype
9, USA: Maryland, Colombia, High Is., ix
(MCZ) [not examined, not traced in MCZ].
[Synonymized by Bram & Bickley, 1963: 6.]
Chrysoperla rufilabris (Burmeister); Garland,
1985a: 93.
Fore wing (Fig. 275) d", 11-12 mm; $,
11.5-12.5 mm. Head marked with broad red
stripe on gena, lateral clypeus and lateral frons;
red post-ocular spot usually present; anterior
margin of vertex sometimes marked with red
stripe, which may extend between antennae.
Maxillary palp black dorsally. Antenna slightly
longer than fore wing. Pronotum marked with
yellow longitudinal median stripe; often with red
spot in anterior corner; prothoracic setae short,
pale. Claw (Fig. 276-278) with large basal dila-
tion, over half length of claw hook (1.85-2.23: 1;
mean 2.02). Fore wing relatively narrow (length:
breadth = 3.1-3.3: 1), apex pointed; costal setae
moderately long (0.8-1.2 mm), inclined towards
wing apex; 5-7 inner gradates, 6-8 outer gra-
dates; gradates black; crossviens black at each
end, except costal crossveins at costa and Psc
crossveins entirely green; cell im long, 1st Rs
crossvein meets im subapically or at apex; 1st Rs
crossvein meets Rs obliquely or at right angles.
Abdomen green with yellow median longitudinal
stripe on tergites and dorso-lateral red spot on
tergites 1 and 2; in some specimens (possibly
showing winter colouration) abdomen dark
brown dorsally, posterior corner of tergite black-
ish brown, pale brown ventrally.
Genitalias (Figs 279-282). Lip of sternite 8+9
broad, rounded; with subapical patch of microse-
tae. Dorsal apodeme of tergite 9 short, extending
a little beyond callus cercus; callus cercus nar-
row, ovate; 24-32 trichobothria. Spinellae
present, numerous small tubercles bearing 6-7
spines. Gonosetae numerous, arranged in lateral
group. Arcessus narrow; dorsal striations absent;
apical membrane long, narrow, pointed at tip,
curved ventrad at 90°; 3^4- mid-lateral microsetae
present or absent. Entoprocessus minute or not
visible, narrow crescentic. Gonarcus with lateral
plates narrow, wedge-shaped; arch narrow. Acu-
men of tignum short, broad, rounded apically.
Genitalia9 (Figs 283-285). Callus cercus
rounded; 27 trichobothria. Subgenitale with
long, narrow basal extension. Ventral impression
of spermatheca variable, shallow or deep; vela
long; duct long.
Remarks. The black gradate crossveins will
serve to distinguish C. rufilabris from all other
North American Chrysoperla species except
some specimens of C. johnsoni and C. downesi.
However, spinellae are absent in the male genita-
lia of C. johnsoni and the gena is often marked
with a brown stripe. In C. rufilabris spinellae are
present in the male genitalia and the gena is
invariably marked red. The long, narrow mem-
branous apex of the arcessus in males of rufila-
bris is unique within the genus.
Geographic variation. Male specimens from
Bermuda, Florida, Louisiana, South Carolina,
Texas and Mexico were found to have 3-4 short
setae situated laterally, mid-way along the arces-
sus. The setae were absent in males examined
from Canada, Florida, Georgia, North Carolina,
Oklahoma, Virginia. Up to 11 such setae were
also reported by Bram & Bickley (1963) in 82%
of specimens examined from Louisiana and 15%
from Maryland, and in specimens from Kansas,
Texas and Florida. Garland (1982) did not find
any Canadian specimens with the setae. I was
unable to find any secondary characters that
conclusively correlated with the presence of the
setae but the majority of the those specimens
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
173
that possessed setae on the arcessus also had long
costal setae and the first Rs crossvein oblique.
The possession of setae on the arcessus is unusual
in Chrysopidae (Brooks & Barnard, 1990) and
would be expected to be of taxonomic signifi-
cance. However, Bram & Bickley (1963) found
that of 12 males reared from a single female, one
had setae on the arcessus and the other 11
specimens had none which suggests this may be
simply a case of intraspecific variation.
Distribution. Canada, Mexico, USA.
Material examined (41cf, 769) Bermuda,
Canada, Mexico, USA: Arkansas, Florida,
Georgia, Louisiana, Massachusetts, Mississippi,
New York, North Carolina, Oklahoma, Oregon,
Pennsylvania, South Carolina, Tennessee,
Texas, Virginia.
SYNOPTIC CHECKLIST OF THE
SPECIES OF CHRYSOPERLA
STEINMANN
(* denotes types examined)
adamsi Henry, Wells & Pupedis, 1993 [Nearctic]
*annae sp. n. [Eastern Palaearctic]
*asoralis (Banks, 1915) [Neotropics)
*oscillans (Navas, 1922) syn. n.
*lindana (Navsis, 1924) syn. n.
*arequipae (Navas, 1929) syn. n.
*carnea (Stephens, 1836) [Palaearctic]
*affinis (Stephens, 1836)
microcephala (Brauer, 1850)
vulgaris (Schneider, 1851)
lamproptera (Stein, 1863)
*lucasina (Lacroix, 1912) syn. n.
angelnina (Navas, 1931)
ferganica (Navas, 1933)
*pictavica (Lacroix, 1933)
adaptata (Navas, 1934) syn. n.
hmdbladi (Tjeder, 1939)
maderensis (Tjeder, 1939) syn. n.
canariensis (Tjeder, 1939)
*shansiensis (Kuwayama, 1962) syn. n.
nanceiensis Semeria, 1980 syn. n.
*comanche (Banks, 1938) [Nearctic]
sperryae (Banks, 1943)
comans (Tjeder, 1966) [Afrotropics]
*manselli Holzel, 1989 syn. n.
*congrua (Walker, 1853) [Old World tropics]
*concolor (Walker, 1853)
otalatis (Banks, 1910) syn. n.
*bequaerti (Navas, 1912)
*lemoulii (Lacroix, 1923) syn. n.
*clypealis (Navas, 1929) syn. n.
solaria (Navas, 1930)
*acutella (Navas, 1933)
*decaryana (Navas, 1934) [Afrotropics]
*defreilasi sp. n. [Neotropics]
downesi (Smith, 1932) [Nearctic]
mohave (Banks. 1938) syn. n.
*dozieri (Smith, 1931) [Haiti]
*exotera (Navas, 1914) [Neotropics]
*extranea (Navas, 1922) syn. n.
estradai (Navas, 1924) syn. n.
*torrei (Navas. 1924) syn. n.
'barbouri (Navas. 1924) syn. n.
*tenera (NavSs. 1924) syn. n.
exterior (Navas. 1925)
cubensis (Navas, 1927)
incisa (Banks, 1948) syn. n.
quadornia (Banks, 1948) syn. n.
castalia (Banks. 1948) syn. n.
'externa (Hagen. 1861) [Neotropics]
"lanata (Banks, 1910)
graciana (Navas, 1919)
*externa cocosensis Adams, 1983 syn. n.
*exut (McLachlan, 1869) [St Helena]
* ' wollastoni (Navas, 1914)
*furcifera (Okamoto, 1914) [Eastern Palaearctic]
savioi (Navas, 1933) syn. n.
*kulingensis (Navas, 1936) syn. n.
galapagoensis (Banks, 1924) [Galapagos Is]
*gallagheri Holzel, 1989 [Oman]
harrisii (Fitch, 1855) [Nearctic]
stenosdgma (Navas, 1914)
*insulata (Fraser, 1957) [Mascarene Is]
*iranica (Holzel. 1967) [Iran]
johnsoni Henry, Wells & Pupedis 1993 [Nearctic]
mediterranea (Holzel, 1972) [Western Palaearctic]
*mexicana sp. n. [Mexico]
*mutata (McLachlan, 1898) [Western Palaearctic]
*nepia (Navas, 1911) syn. n.
*libera (Navas, 1928) syn. n.
*quettana (Navas, 1931) syn. n.
* phaeocephala (Navas, 1931) syn. n.
*grazianii (Navas, 1932) syn. n.
expurgata (Tjeder, 1949)
*concinna (Holzel, 1973) syn. n.
* nigrinervis sp. n. [Canary Is.]
* nipponensis (Okamoto, 1914) stat. rev. (Eastern
Palaearctic]
kurisakiana (Okamoto, 1914)
*ilota (Banks, 1915) syn. n.
174
S.J. BROOKS
*kolthoffi (Navas, 1927) syn. n.
*sinica (Tjeder, 1936) syn. n.
*nyerina (Navas, 1933) [Afrotropics]
*socia (Navas, 1936) syn. n.
*oblita Holzel, 1973 [Eastern Palaearctic]
*plicata (Tjeder, 1966) [Afrotropics]
plorabunda (Fitch, 1855) [Nearctic]
robertsonii (Fitch, 1855)
pseudographa (Fitch, 1855)
illinoiensis (Shimer, 1865)
californica (Coquillet, 1890) syn. n.
*pudica (Navas, 1914) [Afrotropics]
incongrua (Navas, 1914) provisional syn.
*nigriciana (Navas, 1931) syn. n.
*renoni (Lacroix, 1933) [Western Palaearctic]
* ankylopteryformis Monserrat & Diaz-Aranda,
1989 syn. n.
rufilabris (Burmeister, 1839) [Nearctic]
interrupta (Schneider, 1851)
*attenuata (Walker, 1853)
*repleta (Walker, 1853)
novaeboracensis (Fitch, 1855)
tabida (Fitch, 1855)
citri (Ashmead, 1880)
medialis (Banks, 1903)
*siamensis sp. n. [Thailand]
*sillemi (Esben-Petersen, 1935) stat. rev. [Eastern
Palaearctic]
punensis (Ghosh, 1976) syn. n.
gujaratensis (Ghosh, 1976) syn. n.
sanandensis (Ghosh, 1977) syn. n.
zastrowi (Esben-Petersen, 1928) [Afrotropics]
NOT EXAMINED
bellatula Yang & Yang, 1992 (BAU)
chusanina (Navas, 1933) (ACAD SIN)
hainanica Yang & Yang, 1992 (BAU)
longicaudata Yang & Yang, 1992 (BAU)
quinlingensis Yang & Yang, 1989 (BAU)
sola Yang & Yang, 1992 (BAU)
thelephora Yang & Yang, 1989 (BAU)
xizangana (Yang & Yang, 1988) comb. n. (BAU)
yulinica Yang & Yang, 1989 (BAU)
SPECIES NEWLY EXCLUDED FROM CHRYSOP-
ERLA
Mallada anpingensis (Esben-Petersen) comb. n.
Apertochrysa australis (New) comb. n.
Mallada krakatauensis (Tsukaguchi) comb. n.
Mallada maquilingi (Banks) comb. n.
Mallada meloui (Navas) comb. n.
Mallada satilota (Banks) comb. n.
Mallada suzukii (Okamoto) comb. n.
Apertochrysa triactinata (New) comb. n.
REFERENCES
Adams, P.A. 1956. A synonym in the genus Chrysopa (Neu-
roptera: Chrysopidae). Psyche 63: 45.
1959. Neuroptera: Myrmeleontidae and Chrysopidae.
Insects of Micronesia 8 (2): 13-33.
1962. Taxonomy of Hawaiian Chrysopa (Neuroptera:
Chrysopidae). Proceedings of the Hawaiian Entomological
Society 18 (2): 221-223.
1975. Status of the genera Ungla and Mallada Navas
(Neuroptera: Chrysopidae) Psyche 82: 167-173.
1983. A new subspecies of Chrysoperla externa (Hagen)
from Cocos Island. Costa Rica (Neuroptera: Chrysopidae).
Bulletin of the Southern California Academy of Science 82
(1): 42-45.
Adams, P.A. & Penny, N.D. 1987. Neuroptera of the Amazon
Basin. Part 11a. Introduction and Chrysopini. Acta Ama-
zonica 15 (1986): 413-479.
Ashmead, W.H. 1880. Orange insects: a treatise on the injurious
and beneficial insects found on orange trees in Florida.
Jacksonville, Florida, USA. xv + 78 pp.
Aspock, H. 1992. The Neuropteroidea of Europe: a review of
the present knowledge (Insecta). In: Canard, M.. Aspock,
H. & Mansell, M.W. [eds]. Current research in neuropterol-
ogy. Proceedings of the 4th International Symposium on
Neuropterology, 43-56.
Aspock, H., Aspock, U. & Holzel, H. 1980. Die Neuropteren
Europas Vols I & II. Krefeld.
Banks, N. 1903. A revision of the Nearctic Chrysopidae.
Transactions of the American Entomological Society 29:
137-162.
1910a. Some Neuroptera from Australia. Psyche 17 (3):
99-105.
1910b. New South American neuropteroid insects. Pro-
ceedings of the Entomological Society of Washington 12:
146-160.
1915. New neuropteroid insects, native and exotic. Pro-
ceedings of the Academy of Natural Sciences of Philadelphia
66 (1914): 608-632.
1924. Neuroptera from the Williams Galapogos expedi-
tion. Zoologica 5 (17): 177-180.
1938. New Chrysopidae and species new to the United
States. Canadian Entomologist 70: 118-122.
1943. New Neuroptera and Trichoptera from the United
States. Psyche 50 (3-4): 74-81.
1948. Chrysopidae (Nothochrysidae) collected in Mexico
by Dr A. Dampf (Neuroptera). Psyche 55 (4): 151-177.
Barnard, P.C., Brooks, S.J. & Stork, N.E. 1986. The seasonal-
ity and distribution of Neuroptera, Raphidioptera and
Mecoptera on oaks in Richmond Park, Surrey, as revealed
by insecticide knock-down sampling. Journal of Natural
History 20: 1321-1331.
Bickley, W.E. & MacLeod, E.G. 1956. A synopsis of the
nearctic Chrysopidae with a key to the genera (Neuroptera).
Proceedings of the Entomological Society of Washington 58:
177-202.
Bram, R.A. & Bickley, W.E. 1963. The green lacewings of the
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
175
genus Chrysopa in Maryland (Ncuroptcra: Chrysopidae).
Bulletin of the University of Maryland Agricultural Experi-
mental Station A-124: 1 — IS.
Brauer, F. 1850. Bcschreibung und Bcobachtung der osterre-
ichischen Artcn dcr Gaining Chrysopa. Naturwissenschaftli-
cke Abhandlungen Wien 4 14): 1-14.
Brooks, S.J. & Barnard, P.C. 1 WO. The green lacewings of the
world : a generic review (Ncuroptcra: Chrysopidac). Bulletin
of the British Museum (Natural History). Entomology Scries
59(2): 117-286.
Bullini, L. & Cianchi, R. 1984. Electrophoretic studies on
gene-enzyme systems in ehrysopid lacewings. pp. 48-56. In:
Canard. M., Scmeria, Y. and New, T.R. 1984. Biology of
Chrysopidac Junk. The Hague, vii + 294.
Bullini, L., Principi, M.M. & Cianchi, R. 1983 Riccrche
elettroforetiche su specie italianc del generc Chrysopa s.l.
(Ncuroptcra : Chrysopidac). Atti XII Congresso Nazionale
Italiano di Entomologia. Roma. 1980. 235-237.
1984. Electrophoretic studies in the genus Chrysopa (s.l.).
evolutionary and phylogenctic inferences. In: Gepp. J..
Aspock, H. & Holzcl. H. [cds| Progress in the world's
neuropterology. Proceedings of the 1st International Sympo-
sium on Neuropterology . Graz, 1980. 57-59. Graz, Austria.
Burmeister, H.C.C. 1839. Ncuroptcra. In: Hanilhuch der Ento-
mologiel: 757-1050.
Canard, M., Semeria, Y. and New, T.R. 1984. Biology of
Chrysopidac Junk, The Hague, vii + 294.
Canard, M. & Laudeho, Y. 1978. Lcs Ncvropteres captures au
picge dc McPhail dans lcs olivicrs en Grccc. I: L'ilc
d"Aguistri. Biologia Gallo-Hellenical: 65-75.
Cianchi, R. & Bullini, L. 1992 New data on sibling species in
ehrysopid lacewings: the Chrysoperla camea (Stephens) and
Malladu prasinus (Burmeister) complexes (Insecta: Ncu-
roptcra: Chrysopidac): 99-104. In: Canard. M.. Aspock. H.
& Mansell, M.W. 1992. Current research in neuropterology.
Proceedings of the 4th International Symposium on Neu-
ropterology. Toulouse.
Coquillett, D.W. 1890. Report of the California Slate Board of
Horticulture.
Duelli, P. 1987. Eine isolicrte Reliktpopulation von Chrysop-
erla mediterranea (Planipennia: Chrysopidac) in der Sch-
weiz. Mitleilungen der Schweizerischen Enlomologischen
Gesellschaft 60: 301-306.
Esben-Petersen, P. 1928a. Ncuroptcra Planipennia. In: Bei-
trage zur Kenntnis der Land- und SiiSwasserfauna Deutsch-
Siidwestafrikas. Ergcbn. Hamburger deutsch-sudwestafr.
Studienreise 1911,2:203-221.
1928b. Ncuroptcra. Insects of Samoa 7 (3): 89-108.
1935. Myrmeleontidac and Chrysopidac. Karakorum
Zoologie 1: 233-235.
Fitch, A. 1855. Report upon the noxious and other insects of
the State of New York. Transactions of the New York State
Agricultural Society 14: 705-880.
Fraser, F.C. 1957. Odonata and Ncuroptcra of Reunion.
Memoires de ilnslitut Scientifique de Madagascar 8: 15-28.
Garland, J. A. 1982. The taxonomy of the Chrysopidae of
Canada and Alaska. PhD thesis. Department of Entomol-
ogy, McGill Uiversity, Quebec, Canada. 418 pp.
[1985a], Catalogue of Chrysopidae of Canada and Alaska
(Ncuroptera). Neuroptera International 3 (2) (1984): 93-94.
1985b. Identification of Chrysopidae in Canada, with
bionomic notes (Neuroptera). Canadian Entomologist 117:
737-762.
Ghosh, S.K. 1976a. Chrysopa (Chrysoperla) gujuratensis n. sp.
from India (Neuroptera: Chrysopidae). Entomologica Scan-
danavica 7: 74-75.
1976b. A new species of Chrysopa (Neuroptera:
Chrysopidae) from India. Entomon 1 (2): 189-191.
1977. A new species of Chrysopa (Neuroptera: Chrysopi-
dae) from India. Entomon 2 (1): 103-104.
Hagen, H.A. 1861. Synopsis of the Neuroptera of North
America. Smithsonian Institution. Washington. 347 pp.
1866. Hcmcrobidarum Synopsis synonimica. Stetliner
Entomologische Zeilung 27: 369-462
Harris, T.W. 1841. A report on the insects of Massachusetts
injurious to vegetation. 459pp.
Henry, C.S. 1979. Acoustical communication during courtship
and mating in the green laccwing Chrysopa camea (Ncurop.:
Chrysopidac). Annals of the Entomological Society i>l
America 11: 68-79.
1983. Acoustic recognition of sibling species within the
holarctic laccwing Chrysoperla camea (Neuroptera:
Chrysopidae). Systematic Entomology 8: 293-301.
1985a. The proliferation of cryptic species in Chrysoperla
green lacewings through song divergence. Elorida Entomolo-
gist 6S(l): 18-38.
1985b. Sibling species, call differences and speciation in
green lacewings (Neuroptera: Chrysopidac: Chrysoperla).
Evolution 39: 965-984.
1992 [1993|. Chrysoperla mohave (Banks (Ncuroptcra:
Chrysopidae): two familial species in an unexpected dis-
guise Psyche 99 (4): 291-308.
1993. Chrysoperla johnsoni Henry (Neuroptera
Chrysopidac): acoustic evidence lor full species status
Annals of the Entomological Society of America 86 (1):
14-25.
Henry, C.S. & Wells, M.M. 1990. Geographical variation in
the song of Chrysoperla plorahunda in North America
(Neuroptera: Chrysopidae). Annuls of the Entomological
Society of America 83 (3): 317-325
Henry, C.S., Wells, M.M. & Pupedis. R.J. 1993 Hidden
taxonomic diversity within Chrysoperla plorahunda (Ncu-
roptcra : Chrysopidac): two new species based on courtship
SOngS. Annals of the Entomological Society O) America 86 1 1 1:
1-13.
Ilol/cl. H. 1967. Die Ncuroptcrcn Vordcrasicns, BeitrSge zur
Naturkundlichen Forschung in Siidwestdeutschland 26 (1):
19-45.
1970. Zur gencrischen Klassitik.ilion dcr palaarklischen
Chrysopinae. Eine neue Untergattungen der Chrysopidae
(Planipennia). Zeitschrift iter Arbeitsgemeinschaft Osterre-
ichischer Entomologen 22: 44-52.
1972. Anisochrysa (Chrysoperla) mediterranea n. sp. cine
neue europaische Chrysopiden-Spezies (Planipennia.
Chrysopidae). Nachrichienhlalt der Bayensdien Entomolo-
gen 21 (5): 81-83.
1973. Neuroptera aus Nepal. I. Chrysopidae. Khumbu
Hunal 4(3): 333-3XS.
1974. Neue Chrysopidenartcn aus dem Iran (Planipen-
nia). Zeitschrift der Arbgemeinschafi osterreichischer Ento-
mologen 25 (1/2): 41-46.
1980. Insects of Saudi Arabia. Ncuroptera: Fam.
Chrysopidae. Fauna of Saudi Arabia 2:164-173.
1989a. Eine neue Chrysopiden-Specics aus Oman -
Chrysoperla gallaghcri n. sp. (Neuroptera : Chrysopidae).
Entomologische Zeitschrift mil Inscktenborse 99 (5): 58-61.
1989b. Chrysopidae (Neuroptera) der Afrotropischen
region: Genus Chrysoperla Stcinmann. Neuroptera Interna-
tional 5 (3): 165-180.
Holzel, H. & Oh.. P. 1991. Chrysopidac der Mascarene-Inseln
(Neuropteroidea: Planipennia). Entomologische Zeitschrift
101 118): 333-352.
1992. Zoogeographical features of Madagascan Chrysopi-
dae (Insecta : Neuroptera). In: Canard. M., Aspock, H. &
Mansell, M.W. [eds]. Current research in neuropterology.
Proceedings of the 4th International Symposium on Neu-
ropterology. 167-181.
Killington, F.J. 1931. Synonymic notes on certain Neuroptera.
Entomologist 64: 1 12-1 14.
Kimmins, D.E. 1940. Notes on some types of Chrysopidae
176
S.J. BROOKS
(Neuroptera) in the British Museum collections. Annals and
Magazine of Natural History (11)5: 442-449.
Kuwayama, S. 1962. Chrysopidae from Shansi, North China
(Neuroptera). Mushi 36 (2): 9-15.
1966. The type specimens of the Neuroptera in the
collections of the Entomological Institute. Hokkaido Univer-
sity. Insecta Matsumurana 28: 133-140.
Lacroix, J.L. 1912. Faune nevropterique de l'Algerie et de la
Tunisie. Insecta 2: 202-206.
1923. Chrysopides nouveaux. Bulletin de la Societe Ento-
mologique de France 1923: 119-122.
1933. Notes nevropterologiques XI. Lambillionea 33:
146-152.
Leraut, P. 1982. Etude de quelques types de Nevropteres
decrits par J. Lacroix at par L. Navas conserves au Museum
National d'Histoire Naturelle a Paris. Revue Frangaise
d'Entomologie (NS) 4 (1): 48.
1991. Les Chrysoperla de la faune de France (Neur.
Chrysopidae). Entomologie Callica 2 (2): 75-81.
1992. Nevropteres des Alpes centrales francaises (Neur.).
Entomologica Gallica 3 (2): 59-65.
McLachlan, R. 1869. New species, etc. of Hemerobiina: with
synoptic notes (first series). Entomologist's Monthly maga-
zine 6: 21-27.
1898. Neuroptcra-Planipennia collected in Algeria by the
Rev. A.E. Eaton. Transactions of the Entomological Society
of London 1898: 151-168.
Monserrat, V.J. 1977. Neuroptera (Planipennia) de la Sierra
del Guadarrama. Universidad Madrid Catedra de Entomolo-
gia Trabajo 19: 1-202.
Monserrat, V.J. & Diaz-Aranda, L.M. 1989. Nuevos datos
sobre los crisopidos ibericos (Neuroptera. Planipennia:
Chrysopidae). Boletin de la Asociacion Espahola de Entomo-
logia 13: 251-267.
Navas, L. 1908. Neuropteros de Espana y Portugal. Broteria 7:
5-131.
1911. Chrysopides nouveaux (Ins. Neur.). Annates de la
Societe Scientifique de Bruxelles 35: 266-282.
1912. Notes sur quelques Nevropteres d'Afrique. II.
Revue Zoologique Africaine 1: 401-410.
1913. Les chrysopides du Musee de Londres. Annates de
la Societe Scientifique de Bruxelles 37: 292-330.
1914a. Les chrysopides du Musee de Londres. Conclu-
sion. Annates de la Societe Scientifique de Bruxelles 38:
73-114.
1914b. Some Neuroptera from the United States. Bulletin
of the Brooklyn Entomological Society 9: 60-62.
1914c. Neuroptera nova africana. Memorie
dell ' Accademia Ponticia dei Nuovi Lincei 32: 81-108.
1919. Algunos insectos neuropteros de la Repiiblica
Argentina. Revista de la R. Academia de Ciencias Exactas
Fisicas y Naturales de Madrid 17: 287-305.
1922a. Insectos sudamericanos. Revista de la R. Academia
de Ciencias Exactas Fisicas y Naturales de Madrid 19:
255-267.
1922b. Sur des nevropteres nouveaux ou critiques (sixi-
eme serie). Annates de la Societe Scientifique de Bruxelles
42:166-173.
1924a. Algunos insectos de Cuba. Revista de la Real
Academia de Ciencias Exactas, Fisicas y Naturales de Madrid
21 (6): 333-340.
1924b. Crisopidos (Ins. Neur.) neotropicos. Revista Chil-
ena de Historka Natural 27: 1 10-116.
1925. Insectos exoticos nuevos o poco conocidos. Memo-
rias de la Real Academia de Ciencias y Artes de Barcelona 19
(5): 181-200.
1927a. Veinticinco formas nuevas de insectos. Boletin de
la Sociedad Iberica de Ciencias Naturales 26 (9): 48-75.
1927b. Nevropteres de la Chine. Arkiv for Zoologi 19A
(18): 1-5.
1928. Insectos de la Cirenaica. Annali del Museo Civico di
Storia Naturale Ciacomo Doria' 52: 461—464.
1929a. Insecta nova. Series XIII. Neuroptera. Memorie
dell' Accademia Ponticia dei Nuovi Lincei (2) 12: 15-32.
1929b. Insectes du Congo Beige (Series III). Revue de
Zoologie et de Botanique Africaines 18: 92-1 12.
1930. Insecta Nova Series XV. Memorie dell'Accademia
Ponticia dei Nuovi Lincei (2) 14: 409-416.
1931a. Decades de insectos nuevos. Revista de la R.
Academia de Ciencias Exactas Fisicas y Naturales de Madrid
26: 60-86.
1931b. Insectos del Museo de Paris. Broteria 27 (3):
101-136.
1931c. Insectes du Congo Beige. Ser. VI. Revue de
Zoologie et de Botanique Africaines 21: 123-144.
1932. Insetti neurotteri ed affini. Annali del Museo Civico
di Storia Naturale Giacomo Doria' 55: 409-421.
1933a. Neuropteros exoticos. Memorias de la R. Aca-
demia de Ciencias y Artes de Barcelona 23: 203-216.
1933b. Neurotteri e Tricotteri del Deutsches Entomolo-
gisches Institut di Berlino-Dahlcm. Bollettino delta Societa
Entomologica Italiana 65: 105-113.
1933c. Nevropteres et inSectes voisins - Chine et pays
environnants. Notes d'Entomologie Chinoise 1 (9): 1-16.
1934a. Nevropteres et insectes voisins - Chine et pays
environnants. Notes d'Entomologie Chinoise 1 (14): 1-10.
1934b. Insectos de Madagascar. Revista de la Academia de
Ciencias Exactas, Fisico-Quitnicas y Naturales de Zaragoza
17: 49-76.
1936a. Neuroptera. Embioptera, Plecoptcra.
Ephemeroptera et Trichoptera. In: Mission scientifique de
I'Omo. 3. Fasc. 19. Memoires du Museum National
d'Histoire Naturelle (N.5) 4: 101-128.
1936b. Nevropteres et insectes voisins Chine et pays
environnants. Notes d' Entomologie Chinoise 3: 37-62.
Nei, M. 1972. Genetic distance between populations. The
American Naturalist 106: 283-292.
New, T.R. 1980. A revision of the Australian Chrysopidae
(Insecta: Neuroptera). Australian Journal of Zoology Suppl.
77: 1-143.
Ohm, P. & Holzel, H. 1982. Tiergeographische und okolo-
gische Aspekte der Neuropterenfauna der Kapverden. Cou-
rier Forschungsiixstitut Senckenberg 52: 159-165.
1992. Aspects of biogeography and phenology of
Chrysopidae from the Republic of Sudan (Insecta: Neu-
roptera). In: Canard, M., Aspock, H. & Mansell, M.W.
1992. Current research in neuropterology. Proceedings of the
4th International Symposium on Neuropterology, 291-301.
Okamoto, H. 1914. Uber die Chrysopiden-Fauna Japans. Jour-
nal of the College of Agriculture, Tohoku Imperial Univer-
sity, Sapporo, Japan 6: 51-74.
Pongracz, S. 1913. Ujabb adatok Magyarorszag Neuroptera-
faunajahoz. Rovartani Lapok 20: 175-186.
Rambur, M.P. 1842. Histoire naturelle des insectes. Nevrop-
teres. Paris. 534 pp.
Schneider, W.G. 1851. Symbolae as monographiam generis
Chrysopae, Leach. Vratislaviae. 178 pp.
Semeria, Y. 1977. Discussion de la validite taxonomique du
sous-genre Chrysoperla Steinmann. Nouvelle Revue
d' Entomologie 7: 235-238.
1980. Une sous espece nouvelle de Chrysoperla carnea
(Stephens) (Planipennia. Chrysopidae): nanceiensis ssp. nov.
Bulletin de la Societe Entomologique de Mulhouse 1980:
29-30.
1992. Donnees numeriques relatives aux genitalia males
des Chrysoperla Steinmann et leur valeur dans la discrimina-
tion specifique (Insecta: Neuroptera: Chrysopidae). In:
Canard, M., Aspock, H. & Mansell, M.W. [eds]. Current
research in neuropterology. Proceedings of the 4th Interna-
tional Symposium on Neuropterology, 333-339.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
177
Semeria, Y. & Quilici, S. 1986. Premiere contribution a 1'etude
des Chrysopidae (Neuroptcra: Planipennia) de rile de la
Reunion (Ocean Indien). Neuroplera International 4 (2):
107-115.
Shinier, H. 1865. Description of the imago and larva of a new
species oi ( hrysopa, Proceedings of the Entomological Soci-
ety oj Philadelphia 4: 208-212.
Smith, R.C. 1931. The Neuroptcra of Haiti, West Indcs.
Annals of the Entomological Society of America 24: 798-823.
1932. The Chrysopidae (Neuroptcra) of Canada. Annals
of the Entomological Society of America 25: 579-601.
Stein, J.P.E.F. 1863. Bcitrag zur Ncuroptcren-Fauna Griech-
cnlands mit Bcrucksichtigung dalmatinischcr Arten. Berliner
Enlomologische Zeitschrift 7: 4 1 1-422.
Steinmann, H. 1964. The Chrysopa species (Neuroptcra) of
Hungary. Annates Historico-Naturales Musei Nattonales
Hungarici 56: 257-266.
Stephens, J.F. 1836. Illustrations of British entomology. Man-
dibulata 6. London.
Tauber, C.A. 1974. Systematica of North American chrysopid
larvae: Chrysopa cornea group (Neuroptcra). The Canadian
Entomologist 106: 1 133-1153.
Tauber, C.A. & Tauber, M.J. 1973a. Diversification and
secondary intcrgradation of two Chrysopa carnea strains
(Neuroptcra: Chrysopidae). Canadian Entomologist 105 (9(:
1153-1167.
1977a. A genetic model lor sympatric speciation through
habitat diversification and seasonal isolation. Nature 268:
702-705.
1977b. Sympatric speciation based on allelic changes at
three loci: evidence from natural populations in two habitats.
Science 197: 1298-1300.
1982a. Sympatric speciation in Chrysopa. further discus-
sion. Annals of the Entomological Society oj America 75:
1-2.
1982b. Evolution of seasonal adaptations and life history
traits in Chrysopa. response to diverse selective pressures, pp
51-72. In: Dingle, H. & Hegmann, J. P. |cds| Evolution and
genetics of life histories. Springer- Verlag, New York.
1985. Eeophysiologieal responses in life-history evolution:
evidence for their importance in a geographically widespread
insect species complex. Canadian Journal of Zoology 64:
875-884
1987. Inheritance of seasonal cycles in Chrysoperla
(Insecta: Neuroptcra). Genetic Research 49: 215-223.
1973b. Nutritional and photoperiodic control of the
seasonal reporductive cycle in Chrysopa mohave (Neu-
roptera). Journal of Insect Physiology 19: 729-736.
1981 . Seasonal responses and their geographic variation in
Chrysopa dotvnesi: eeophysiologieal and evolutionary con-
siderations. Canadian Journal of Zoology 59 (3): 370-376.
Thierry, D. & Adams, P.A. 1992. Round table discussion on
the Chrysoperla carnea (Stephens) complex (Insecta: Neu-
roptcra: Chrysopidae). ///: Canard, M., Aspock, H. &
Mansell, M.W. 1992. Current research in ncuropterology.
Proceedings of the 4th International Symposium on Neu-
ropterology . 367-373.
Thierry, D., Cloupeau, R. & Jarry, M. 1992. La chrysope
commune Chrysoperla carnea (Stephens) sensu lato dans le
centre de la France: mise en evidence dun complexe
d'especes (Insecta: Neuroptcra: Chrysopidae): 379-392 In:
Canard, M.. Aspock, H. & Mansell, M.W. 1992. Current
research in ncuropterology. Proceedings of the 4th Interna-
tional Symposium on Neuropterology. 379-392.
Tjeder, B. 1936. Schwcdish-chinesischc wissenschaftliche
Expedition nach den nordwcstlichcn Provinzcn Chinas.
Arkivfor Zoologi 29A (8): 1-36.
1939. Die Arthropodcnfauna von Madeira nach den
Ergebnissen der Reise von Prof. Dr. O. Lundblad, Juli-Au-
gusl 1935. XVI. Neuroptcra. Arkiv for Zoologi 31A (Nr. 15):
58pp.
1949. Two new Chrysopidae from Palestine (Ncur.).
Opuscula Entomologica 14: 81-84
1966. Neuroptcra - Planipennia 5. Family Chrysopidae.
South African Animal Life 12: 228-534
1971. Rcdescriptions and lectotype designations of some
exotic Chrysopidae (Ncur.). Entomologica Scandanayica 2:
110-188.
Tsukaguchi. S. 1985. A checklist of published species of
Japanese Chrysopidae (Neuroptcra). Kontyu 53: 503-506.
Walker, F. 1853 list oj the specimens of neuropterous insects in
the collections of the British Museum. II. Sialides-
Nemoplerides : 193-476.
Wells, M.M. 1991. Reproductive isolation and genetic diver-
gence among populations of green lacewmgs of the genus
Chrysoperla. PhD Thesis, The University of Connecticut
115pp.
Yang, C.-k. 1988. Neuroptcra: Osmylidac. Dilandae. llemero-
biidae, Chrysopidae, Mantispidae, Myrmelcontidae, Ascal-
aphidae. Corydalidac. Insects of Mt Namjugharwu region oj
Xizang. 193-212.
Yang, C.-k. & Yang, X.-k. 1989. Fourteen new species of green
lacewings from Shaanxi Province (Neuroptcra: Chrysopi-
dae). Entomotaxonomia 11 (1-2): 13-30.
Yang, X.-k. & Yang, C.-k. 1990a. Examinations and rcdescrip-
tions of the type specimens of some Chinese Chrysopidae
(Neuroptcra) described by L. Navas. Neuroptcra Interna-
tional 6 (2): 75-83.
Yang, C.-k. & Yang, X.-k. 1990b. A revision of the Chinese
Malluda (Neuroptcra: Chrysopidae). Scientific treatise on
Systematic and Evolutionary Zoology 1: 135-149.
Yang, X.-k. & Yang, C.-k. 1992 Study on the genus Chrysop-
erla (Neuroptcra: Chrysopidae). Acta Entomologica Sinica
35(1): 78-86.
Zimmerman, E.C. 1957. Insects of Hawaii. Volume 6.
Ephemeropleru - Neuroptcra - I richoptera. University of
Hawaii, Honolulu. 209 pp.
178
SJ. BROOKS
Figs 1-13 1-4, 6-13, Chrysoperla carnea; 5, C. sillemi. 1, fore wing; 2, fore wing f. typica (London, UK); 3, fore
wing f. lucasina (Hertfordshire, UK); 4, hind tarsal claw (Spain); 5, hind tarsal claw (Pakistan); 6, hind tarsal
claw (Crete); 7, hind tarsal claw (Israel); 8, apex d" abdomen lateral; 9, 0" gonarcus complex, dorsal; 10, C?
gonarcus complex, lateral; 11, apex $ abdomen, lateral; 12, $ subgenitale, caudal; 13, $ spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
179
15
Figs 14-23 14. 16, Chrysoperla downesi; 15, 21-22 C. plorabunda; 17, 19-20 C. carnea; 18, C. sillemi; 23, C.
johnsoni. 14, fore wing; 15, hind tarsal claw; 16 hind tarsal claw; 17, O" tignum (Iran); 18, d* tignum (India); 19,
O" tignum f. typica (London, UK); 20, C? tignum f. lucasina (Hertfordshire, UK); 21, cf tignum (Canada); 22,
O" tignum (Tuscon, USA); 23 hind tarsal claw.
180
S.J. BROOKS
Figs 24-31 Chrysoperla zastrowi. 24, fore wing; 25, hind tarsal claw, lateral; 26, apex c? abdomen, lateral; 27, cf
gonarcus complex, dorsal (gonosaccus not shown); 28, d" tignum, dorsal; 29, apex 9 abdomen, lateral; 30, 9
subgenitale, caudal; 31, $ spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
181
Figs 32-39 Chrysoperla furcifera. 32, fore wing; 33, hind tarsal claw, lateral; 34, apex cf abdomen, lateral; 35, d*
gonarcus complex, dorsal; 36, cf tignum, dorsal; 37, apex $ abdomen, lateral; 38, 9 subgenitale, caudal; 39,' $
spermatheca, lateral.
182
S.J. BROOKS
Figs 40-49 Chrysoperla iranica. 40, fore wing; 41. hind tarsal claw, lateral; 42, apex cf abdomen, lateral; 43,
sternite 8+9, ventral; 44, a" tignum, dorsal; 45, cf gonarcus complex, lateral; 46, a" gonarcus complex, dorsal
(gonosaccus not shown); 47, apex $ abdomen, lateral; 48, $ subgenitale, caudal; 49, $ spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
183
Figs 50-58 Chrysoperla mediterranea. 50, fore wing; 51, hind tarsal claw, lateral; 52, apex o" abdomen, lateral;
53, cf tignum, dorsal; 54, cf gonarcus complex, dorsal; 55, O" gonarcus complex, lateral; 56, apex 9 abdomen,
lateral; 57, $ subgenitale, caudal; 58. 9 spermatheca, lateral.
184
S.J. BROOKS
Figs 59-67 Chrysoperla nipponensis. 59, fore wing; 60, hind tarsal claw, lateral (Japan); 61, hind tarsal claw,
lateral (Mongolia); 62, apex cf abdomen, lateral; 63, cf tignum, dorsal; 64, cf gonarcus complex, dorsal; 65,
apex $ abdomen, lateral; 66, $ subgenitale, caudal; 67, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
185
Figs 68-75 Chrysoperla renoni. 68, fore wing; 69, hind tarsal claw, lateral; 70, apex O" abdomen, lateral; 71, c?
gonarcus complex, dorsal; 72, cf tignum, dorsal; 73, apex 9 abdomen, lateral; 74, $ subgenitale, caudal; 75, $
spermatheca, lateral.
186
S.J. BROOKS
Figs 76-83 Chrysoperla oblita. 76, fore wing; 77, hind tarsal claw, lateral; 78, apex d" abdomen, lateral; 79, ($
gonarcus complex, dorsal; 80, cf tignum, dorsal; 81, apex 9 abdomen, lateral; 82, 9 subgenitale, caudal; 83, 9
spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
187
Figs 84-89 Chrysoperla annae. 84, fore wing; 85, hind tarsal claw, lateral; 86, apex a" abdomen, lateral; 87, d"
tignum, dorsal; 88, d* gonarcus complex, lateral; 89, d" gonarcus complex, dorsal.
188
S.J. BROOKS
Figs 90-98 Chrysoperla gallagheri. 90, fore wing; 91, hind tarsal claw, lateral; 92, apex d" abdomen, lateral; 93,
Cf gonarcus complex, dorsal (gonosaccus not shown); 94, cf gonarcus complex, lateral; 95, cf tignum, dorsal;
96, apex $ abdomen, lateral; 97, 9 subgenitale, caudal; 98, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
189
100 x^II^
Figs 99-108 Chrysoperla comans. 99, fore wing; 100, hind tarsal claw, lateral; 101, apex (f abdomen, lateral;
102, sternite 8+9 of cf abdomen, ventral; 103, cf gonarcus complex, lateral; 104, cT gonarcus complex, dorsal;
105, cf tignum, dorsal; 106, apex 9 abdomen, lateral; 107, $ spermatheca, lateral; 108, 9 subgenitale, caudal.
190
S.J. BROOKS
Figs 109-114 Chrysoperla decaryana. 109, fore wing; 110, hind tarsal claw, lateral; 111, apex c? abdomen,
lateral; 112, cf tignum, dorsal; 113, ($ gonarcus complex, dorsal; 114, cf gonarcus complex, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
191
Figs 115-123 Chrysoperla siamensis. 115, fore wing; 116, hind tarsal claw, lateral; 117, apex cS abdomen, lateral;
118, cf gonarcus complex, lateral; 119, o" gonarcus complex, dorsal; 120, cf tignum, dorsal; 121, apex 9
abdomen, lateral; 122, $ spermatheca, lateral; 123, 9 subgenitale, caudal (basal extension shown folded
underneath subgenitale).
192
S.J. BROOKS
Figs 124-132 Chrysoperla nyerina. 124, fore wing; 125, hind tarsal claw, lateral; 126, apex cf abdomen, lateral;
127, cf tignum, dorsal; 128, cf gonarcus complex, dorsal; 129, cf gonarcus complex, lateral; 130, apex $
abdomen, lateral; 131, $ subgenitale, caudal; 132, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
193
Figs 133-141 Chrysoperla plicata. 133, fore wing; 134, hind tarsal claw, lateral; 135, apex d" abdomen, lateral;
136, cf tignum, dorsal; 137, ($ gonarcus complex, dorsal; 138, a" gonarcus complex, lateral; 139, apex 9
abdomen, lateral; 140, 9 subgenitale, caudal; 141, 9 spermatheca, lateral.
194
S.J. BROOKS
Figs 142-148 Chrysoperla insulata. 142, fore wing; 143, hind tarsal claw, lateral; 144, apex c? abdomen, lateral;
145, d" gonarcus complex, lateral; 146, apex 9 abdomen, lateral; 147, 9 subgenitale, caudal; 148, 9
spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
195
Figs 149-159 Chrysoperla pudica. 149, fore wing; 150, hind tarsal claw, lateral; 152, apex cf abdomen, lateral;
153, cf tignum, dorsal (Transvaal); 154, cf tignum, dorsal (Nigeria); 155, cf gonarcus complex, lateral; 156, cf
gonarcus complex, dorsal; 157, apex 9 abdomen, lateral; 158, $ subgenitale, caudal; 159, $ spermatheca,
lateral.
196
S.J. BROOKS
Figs 160-170 Chrysoperla mutata. 160, fore wing; 161, hind tarsal claw, lateral (Punjab); 162, hind tarsal claw,
lateral (Arabia); 163, hind tarsal claw, lateral (Algeria); 164, apex cf abdomen, lateral; 165, spinellae; 166, cf
gonarcus complex, lateral; 167 cf tignum, dorsal; 168, apex £ abdomen, lateral; 169, $ subgenitale, caudal;
170, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
197
Figs 171-178 Chrysoperla nigrinervis. 171, fore wing; 172, apex d" abdomen, lateral; 173, cf tignum, dorsal; 174,
Cf gonarcus complex, lateral; 175, apex 9 abdomen, lateral; 176, $ subgenitale, caudal; 177, 9 spermatheca,
lateral; 178 hind tarsal claw, lateral.
198
S.J. BROOKS
Figs 179-187 Chrysoperla exul. 179, fore wing; 180, hind tarsal claw, lateral; 181, apex cf abdomen, lateral; 182,
Cf gonarcus complex, lateral; 183, cf gonarcus complex, dorsal; 184, cf tignum, dorsal; 185, apex $ abdomen,
lateral; 186, $ subgenitale, caudal; 187, $ spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
199
Figs 188-195 Chrysoperla congrua. 188, fore wing; 189, hind tarsal claw, lateral; 190, apex O" abdomen, lateral;
191, cf tignum, dorsal; 192, cf gonarcus complex, dorsal; 193, apex 9 abdomen, lateral; 194, 9 subgenitale,
caudal; 195, 9 spermatheca, lateral.
200
S.J. BROOKS
Figs 196-202 Chrysoperla dozieri. 196, fore wing; 197, hind tarsal claw, lateral; 198, apex o" abdomen, lateral;
199, cf tignum, dorsal; 200, d" apex of arcessus; 201, cf gonarcus complex, lateral (gonosaccus not shown); 202,
gonarcus complex, dorsal (gonosaccus not shown).
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
201
205
Figs 203-208 Chrysoperla defreitasi. 203, fore wing; 204, apex cf abdomen, lateral; 205, d" gonarcus complex,
dorsal; 206, cf gonarcus complex, lateral; 207, cf tignum, dorsal; 208, hind tarsal claw, lateral.
202
S.J. BROOKS
209
214
Figs 209-217 Chrysoperla asoralis. 209, fore wing; 210, hind tarsal claw, lateral; 211, apex 9 abdomen, lateral;
212, 9 spermatheca, lateral; 213, $ subgenitale, caudal; 214, apex cf abdomen, lateral; 215, cf tignum, dorsal;
216, cf gonarcus complex, dorsal; 217, cf gonarcus complex, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
203
219
999
Figs 218-226 Chrysoperla exotera. 218, fore wing; 219, hind tarsal claw, lateral; 220, apex o" abdomen, lateral;
221, cT gonarcus complex, dorsal; 222, O" gonarcus complex, lateral; 223, d" tignum, dorsal; 224, apex $
abdomen, lateral; 225, 9 subgenitale, caudal; 226, 9 spermatheca, lateral.
204
S.J. BROOKS
229
Figs 227-236 Chrysoperla externa. 227, fore wing; 228, hind tarsal claw, lateral; 229, hind tarsal claw, lateral;
230, apex d" abdomen, lateral; 231, c? tignum, dorsal; 232, o" gonarcus complex, dorsal; 233, d" apex of
arcessus, dorsal; 234, apex 9 abdomen, lateral; 235, $ subgenitale, caudal; 236, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
205
Figs 237-245 Chrysoperla galapagoensis. 237, fore wing; 238, hind tarsal claw, lateral; 239, apex d" abdomen,
lateral; 240, ($ gonarcus complex, caudal; 241, cf tignum, dorsal; 242, apex 9 abdomen, lateral; 243, 9
subgenitale, caudal; 244, 9 subgenitale, ventral; 245, 9 spermatheca, lateral.
206
S.J. BROOKS
Figs 246-255 Chrysoperla comanche. 246, fore wing; 247, hind tarsal claw, lateral; 248, apex cf abdomen,
lateral; 249, cf tignum, dorsal; 250, cf gonarcus complex, dorsal; 251, cf gonarcus complex, lateral; 252, apex
of arcessus; 253. apex $ abdomen, lateral; 254. 9 subgenitale, caudal; 255, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
207
256
260
Vy 265
Figs 256-265 Chrysoperla harrisii. 256, fore wing; 257, hind tarsal claw, lateral; 258, apex cf abdomen, lateral;
259, cf gonarcus complex, dorsal; 260, cf apex of arcessus; 261, cf gonarcus complex, lateral; 262, cf tignum,
dorsal; 263, apex $ abdomen, lateral; 264, 9 spermatheca, lateral; 265, 9 subgenitale, caudal.
208
S.J. BROOKS
266
Figs 266-274 Chrysoperla mexicana. 266, fore wing; 267, hind tarsal claw, lateral; 268, apex cf abdomen,
lateral; 269, cf tignum, dorsal; 270, cf gonarcus complex, dorsal; 271, cf apex of arcessus; 272, apex 9
abdomen, lateral; 273, $ subgenitale, caudal; 274, 9 spermatheca, lateral.
TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA
209
Figs 275-285 Chrysoperla rufilabris. 275, fore wing; 276, hind tarsal claw, lateral (Texas, USA); 277, hind tarsal
claw, lateral (Canada); 278, hind tarsal claw, lateral (North Carolina, USA); 279, apex (S abdomen, lateral;
280, cf tignum, dorsal; 281, cf apex of arcessus; 282, o" gonarcus complex, dorsal (gonosaccus not shown); 283,
apex 9 abdomen, lateral; 284, 9 spermatheca, lateral; 285, 9 subgenitale, caudal.
210
S.J. BROOKS
INDEX
This index is to species-group and genus-group names only. Principal references are in bold; synonyms are in italics. References to
the key (p. 144) and the checklist (p. 173) are not included.
acutella 165
adamsi 142, 150
adaptata 146. 147
affinis 146
Anachrysa 141
angelnina 146
ankylopteryformis 156
annae 142, 157
Anomalochrysa 141
anpingensis (Mallada ) 144
Apertochrysa 141, 142, 144
arequipae 167
asoralis 143. 167
Atlantochrysa 141, 142
attenuata 172
australis (Apertochrysa) 144
barbouri 167
bellatula 144
bequaerti 164
boninensis (Mallada) 144
brevicollis 161, 162
calif or nica 150
canariensis 146
carnea 142, 146, 149, 152, 154, 156, 157
castalia 168
Chrysopidia 141
Chrysotropia 141
chusanina 155
citri 172
clypealis 164
cocosensis 168, 169
comanche 143, 168, 169, 170, 171
comans 142, 158, 159
concinna 163
concolor 164
congrua 143, 164, 165, 169
cubensis 167
Cunctochrysa 141
decaryana 142, 158, 159
defreitasi 143, 166
desjardinsi (Mallada) 144
Dichochrysa 141, 142, 144
downesi 142, 151, 152, 172
dozieri 143, 165, 166
estradai 167
exotera 143, 166, 167, 170
expurgata 163
exterior 167
externa 143, 166, 168, 169, 170
extranea 167
exul 143, 164
ferganica 146
flava (Nineta) 147
furcifera 142, 146, 152, 153
galapagoensis 143. 169
gallagheri 142, 157, 159
graciana 168
grazianii 163
gujaratensis 149
hainanica 144
harrisii 143, 170, 171
illinoiensis 150
ilota 155
incisa 167
incongrua 161. 162
inconspicua (Mallada) 144
insulata 142. 161
interrupla 171
iranica 142, 153
italotis (Plesiochrysa) 165
johnsoni 142. 150, 172
kolthoffi 148. 155. 156
krakatauensis (Mallada) 144
kulingensis 152
kurisakiana 155
lamproplera 146
lanata 168
lemouhi 164
libera 162
lindana 167
longicaudata 144
lucasina 146, 148
lundbladi 146
maderensis 146
Mallada 141, 144
manselli 158
maquilingi (Mallada) 144
medial is 172
mediterranea 142, 146, 154, 156
Meleoma 142
meloui (Mallada) 144
mexicana 143, 171
microcephala 146
mohave 150, 151. 152
mutata 143. 158, 162, 163, 164
nanceiensis 146
nepia 162
nigriciana 161
nigrinervis 143, 163, 164
nipponensis 142, 146. 155
novaeboracensis 172
nyerina 142. 158. 160. 161
oblita 142. 146, 156, 157
occipitalis 165
oscillans 167
otalatis 164
Pcyerimhoffina 141
phaeocephala 162
pictavica 146
plicata 142. 158. 160, 161
plorabunda 142. 146, 150, 151
pseudographa 150
pudica 143. 158, 161, 162, 163
punensis 149
quadornia 167
queitana 162
quinlingensis 144
renoni 142, 156
repleta 172
robertsonii 150
rubella 165
rufilabris 143. 167. 171. 172
sanandensis 149
satilota (Mallada) 144
savioi 152, 153
shansiensis 146, 147
siamensis 142, 159
sillemi 142. 149
sinica 155
socia 160
sola 144
solaria 165
sperryae 170
slenostigma 170
Suarius 142
suzukii (Mallada) 144
tabida 172
tenera 167
thelephora 144
torrei 167
triactinata (Apertochrysa) 144
vittata (Nineta) 147
vulgaris 146
wollastoni 164
xizangana 144
yulinica 144
zastrowi 142, 146, 149, 152
Bulletin of The Natural History Museum
Entomology Series
Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside
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Volume 44
No. 1 The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Diptera:
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Pp. 1-30, 96 figs, 2 tables. £4.50
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Volume 47
No. 1 A new genus of oriental lacewings (Neuroptera: Chrysopidae). S.J. Brooks. 1983. Pp. 1-26,
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No. 1 Afrotropical jumping plant lice of the family Triozidae (Homoptera: Psylloidea-. D. Hollis. 1984.
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No. 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). P.S. Broomfield.
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Volume 51
No. 1 The ichneumon-fly genus Banchus (Hymenoptera) in the Old World. M.G. Fitton. 1985.
Pp. 1-60, 129 figs. £10.80
No. 2 The phylogeny, classification and evolution of parasitic wasps of the subfamily Ophioninae
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No. 1 The sandflies of Egypt (Diptera: Phlebotominae). R.P. Lane. 1986. Pp. 1-35, 80 figs,
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No. 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). G.S. Robinson. 1986.
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No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986.
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No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera.
J. Quinlan. 1986. Pp. 243-366, 359 figs. £21.00
Volume 53
No. 1 A review of Miletini (Lepidoptera: Lycaenidae). J.N. Eliot. 1986. Pp. 1-105, 108 figs. £18.00
No. 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini. I.D. Gauld & G.A.
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No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their
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No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble.
1986. Pp. 251-286, 102 figs. £7.00
Volume 54
No. 1 Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae). S. Okajima.
1987. Pp. 1-74, 207 figs. £14.00
No. 2 Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae
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No. 1 A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of
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Volume 56
No. 1 The legume-feeding psyllids (Homoptera) of the wet Palaearctic Region. I.D. Hodkinson & D.
Hollis. 1987. Pp. 1-86, 294 figs. £16.00
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Pp. 129-170, 59 figs. £8.00
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Pp. 171-229, 199 figs. £11.00
Volume 57
No. 1 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with
special reference to the fauna of Costa Rica. I.D. Gauld. 1988. Pp. 1-309, 352 figs,
32 maps. £52.00
No. 2 A taxonomic revision of Alabagrus (Hymenoptera: Braconidae). M.J. Sharkey. 1988.
Pp. 311-437, 28 figs, 22 maps. £24.50
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221 figs. £25.00
Volume 58
No. 1 The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78.
40 figs.
No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M.
Arzube & C.A. Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour).
Volume 59
No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their
taxonomy (Orthoptera: Tettigoniidae). DR. Ragge. 1990. Pp. 1-35.
A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek.
1990. Pp. 37-115.
No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks &
P.C. Barnard. 1990. Pp. 117-286.
Volume 60
No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). PH. Williams.
1991. Pp. 1-204.
No. 2 Sattkria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M.
Pitkin & K. Sattler. 1991. Pp. 205-241.
A review of wing reduction in Lepidoptera. K. Sattler. 1991 . Pp. 243-288.
Volume 61
No. 1 Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76.
No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx
Warren (Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148.
Volume 62
No. 1 Caloptilia leaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S.
Robinson. 1993. Pp. 1-37.
No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular
reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M.
Pitkin. 1993. Pp. 39-159.
Volume 63
No. 1 A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of
the immature stages of economically important insect species (mainly Hemiptera and
Lepidoptera). D.W. Huang and J. S. Noyes. Pp. 1-135.
'-■■
137 A taxonomic review of the common green lacewing genus Chrysoperla
(Neuroptera: Chrysopidae)
S.J. Brooks
Natural his i oRY
MUSEUM SHOPS
139 -5^
ENTOMOLOGY SERIES
Vol. 63, No. 2, November 1994
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